Seasonal dynamics of forest spiders (Arachnida: Araneae) in the

Munibe (Ciencias Naturales-Natur Zientziak) • NO. 57 (2009) • 83-146 • ISSN 0214-7688
Seasonal dynamics of forest spiders (Arachnida:
Araneae) in the temperate zone of the Basque
Country and Navarra (northern Spain)
A. CASTRO GIL 1
ABSTRACT
The aims of this research are to determine the time of year with the highest species richness to avoid whole-year samplings in future studies, and to contribute to
the seasonal dynamics and faunistic knowledge of the spider fauna of the study area.
Hence, it summarizes temporal changes in species richness and seasonal activities
of spiders collected in several temperate forests sampled using different methods.
Results show the May-June transition as the period with the peak in species richness. However, this maximum takes place earlier in the epigeal stratum (May), and
later on tree trunks (June). Additionally, lower strata show higher representation of
species of long mating periods. Three species, Centromerita bicolor, Micrargus apertus and Midia midas are new records for Iberian fauna, while Peponocranium ludicrum is a new genus record for Spain. Seasonal activities of Nemesia simoni and
Labulla flahaulti are described for the first time. Data obtained in two of the sampled forests will make it possible to evaluate the efficiency of future short-term
collecting protocols for the study area.
• KEY WORDS: Araneae, seasonal activity, temperate forests, species richness, Spain.
RESUMEN
Los objetivos del presente trabajo son determinar la época del año que registra el
mayor número de especies a fin de evitar muestreos de ciclos anuales completos en
estudios futuros y contribuir al conocimiento faunístico y de las dinámicas estacionales de la araneofauna del área de estudio. Para ello, se compendia las variaciones
temporales de la riqueza específica y de las actividades estacionales de las arañas
capturadas en varios bosques templados muestreados a través de distintos métodos.
Los resultados determinan la transición entre Mayo y Junio como el periodo que presenta el máximo de riqueza específica. Sin embargo, este máximo tiene lugar antes
en el estrato epigeo (Mayo) y más tarde (Junio) en los troncos de los árboles.
Además, los estratos más bajos muestran una mayor representación de especies de
periodos de reproducción largos. 3 nuevas especies, Centromerita bicolor,
Micrargus apertus y Midia midas, se citan para la Península Ibérica y un nuevo
género, Peponocranium ludicrum, para España. Las actividades estacionales de
1 Sociedad de Ciencias Aranzadi / Zientzia Elkartea. Departamento de Entomología.
Zorroagagaina 11 • 21014 Donostia - San Sebastián.
83
A. CASTRO GIL
Nemesia simoni y Labulla flahaulti se describen por primera vez. Los datos obtenidos en dos de los bosques muestrados permitirán evaluar la eficacia de futuros protocolos de recolección a corto plazo para el área de estudio.
• PALABRAS CLAVE: Araneae, actividad estacional, bosques templados, riqueza específica, España.
LABURPENA
Lan honen helburua, aztertzen den eremuan armiarma-fauna espezie kopuru handiena urtearen zein sasoietan ematen den zehaztea da, hurrengo azterketetan urteetako ziklo osoetan laginketa neketsuak egin beharra ekiditeko, ezagutza faunistikoan eta urte-sasoien zehar ematen den dinamika ezagutzen laguntzearekin batera.
Horretarako, metodo desberdinak erabiliz lagindu diren hainbat baso epeletan
harrapatutako armiarmak aurkezten duten aberastasun espezifikoa eta urte-sasoietan
erakusten dituzten jarduerak aurkezten dira. Emaitzen arabera, maiatzetik ekainerako bitartea da aberastasun espezifiko handieneko epea. Dena dela, gehiengo hau
geruza epigeoan lehenago ematen da (maiatza) eta beranduago (ekaina) zuhaitzen
enborretan. Gainera, beherago dauden geruzak, ugalketa epealdi luzeagoak dituzten espezie gehiago dituzte. Iberiar Penintsularako hiru espezie berri, Centromerita
bicolor, Micrargus apertus eta Midia midas, eta, Espainiarako, genero berri bat,
Peponocranium ludicrum, aipatzen dira. Nemesia simoni eta Labulla flahaulti-k
urte-sasoietan dituzten jarduerak lehendabizikoz deskribatzen dira. Lagindu direneko bi basoetan lortutako datuei esker, epe motzean eta azterketa eremurako, etorkizuneko bilketarako protokoloen eraginkortasuna neurtu ahal izango dugu.
• GAKO-HITZAK: Araneae, urte-sasoietako jarduera, baso epelak, aberastasun espezifikoa, Espainia.
wy
INTRODUCTION
Registering the entire biological diversity at regional scale is an impossible task,
whatever economic and time resources are available, since biological diversity is
constantly changing. The use of surrogate taxa as biodiversity indicators of a specific area has been proposed as a way of addressing this problem and facilitating biodiversity management and conservation policies (WILLIAMS et al., 1997; VANCLAY,
2004). Spiders are often used as biodiversity indicators for the following reasons
(MARC et al., 1999; PLATNICK, 1999; MAELFAIT et al., 2004; PEARCE & VENIER,
2006): they show high species richness, play an important role in terrestrial ecological webs, are of economic interest, are easy and inexpensive to sample (yielding
quantitative analyzable results), their taxonomy is reasonably well known in developed countries, and their communities are sensitive to environmental change.
Knowledge of spider fauna is far from being complete both in the Iberian
Peninsula (MORANO, 2004; CARDOSO, 2008) and in the area covered by this study
(CASTRO, 2004a). To counter this lack of information, maximizing economic and
84
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
time resources, intensive short-time sampling protocols have been developed in
other countries (CODDINGTON et al., 1991, 1996; SORENSEN et al., 2002; SCHARFF
et al., 2003; CARDOSO et al., 2008). The first step in applying these methods is to
conduct sampling programs encompassing entire year cycles to ascertain at what
time of year the highest spider diversity is found (CARDOSO, 2004, CARDOSO et
al., 2007).
Potentially, almost the entire surface of the study area would be taken up by
forest ecosystems (ASEGINOLAZA et al., 1996). Forestry covers 61% of the Basque
Country’s temperate area, although more than half of the forest surface is occupied
by plantations of exotic species (DEPARTAMENTO DE AGRICULTURA, PESCA Y
ALIMENTACIÓN 2007). The situation in the temperate zone of Navarra is similar,
though native species predominate (ICONA, 1994). In any case, most indigenous
forests are exploited for sylvicultural and shepherd activities (LOIDI & BASCONES,
1995; ASEGINOLAZA et al., 1996). In this regard, knowledge of spider communities
in temperate forests can be applied for designing suitable biodiversity conservation
programs in the study area, as is happening in other European regions (RIECKEN,
1998; DE BAKKER et al., 2002).
This study therefore analyses collections of woodland spiders available for the
study area with the following objectives: 1) To determine which period of the year
would be most suitable for intensive short-time sampling programs yielding a representative range of regional spider fauna, and 2) to contribute to an understanding
of seasonal dynamics and a faunistic knowledge of spider fauna in the study area.
STUDY
AREA
The data came from four independent studies whose locations are summarized in
Table I. All them show oceanic temperate climate (RIVAS-MARTINEZ, 1994). The
characteristics of the different forest sites sampled are described below:
Site
Gorbea
Artikutza
Town/Province
U.T.M.
Coordinates
Altitude
(m)
Orientation
Termotipe/
Ombrotipe
Areatza-Villaro/Bizkaia
30TWN1673
400-510
Diverse
C/Hi
Goizueta/Navarra
30TWN9786
575-650
SW
C/U
30TWN4070
30TWN6760
30TWN5591
30TWN4990
30TWN6194
30TWN8464
450
390
310
225
20
312
E
W
E
SW
SW
SE
C/Hi
C/Hi
C/Hi
C/Hi
T/H
C/Hi
50-70
E
T/Hi
Arrasate/Gipuzkoa
Ataun/Gipuzkoa
Evergreen Deba-Itziar/Gipuzkoa
oak forests Mendaro/Gipuzkoa
Zumaia/Gipuzkoa
Larraun/Navarra
Igara
San Sebastián/Gipuzkoa 30TWN7894
Table I.- Location and climate of the studied sites. Abbreviations: C = coline, T = thermocoline,
Hi = lower hyperhumid, U = ultrahyperhumid, H = upper humid.
85
A. CASTRO GIL
Gorbea: Samples were taken in the area of Upomakatxa, from seven forest stands
near to each other: five plantations of Pinus radiata D. Don (three 30 year-old plantations, one 18-year-old and one 8-year-old), and two indigenous stands—one small
beech forest (Fagus sylvatica L.) and one young deciduous broadleaf mixed forest
(with predominance of Quercus robur L., Castanea sativa Miller and Corylus avellana L.). The main objective of this study was to analyse the impact of exotic pine
forest plantations on soil invertebrate fauna. Livestock was commonly present in all
stands. More details on the sampling design and first results obtained are given in
BARRAQUETA (1985, 1988, 2001).
Artikutza: Samples were taken from a 5 ha interface between a beech forest and
a deciduous broadleaf forest (with predominance of Quercus robur L.). This stand
was in a regeneration process after a plantation of Scots pine (Pinus sylvestris L.)
had lain abandoned for 70 years. The forest contained a large amount of dead
wood. Livestock was also present at this site. The original purpose of this research
was to study the spatial activity patterns of several families of Hymenoptera
(MARTÍNEZ DE MURGUÍA, 2002). More details of the study area may be found in
MARTÍNEZ DE MURGUÍA et al. (2001, 2002).
Cantabrian evergreen oak forests: This sampling program was carried out in the
six most representative Cantabrian evergreen oak forests in the provinces of
Gipuzkoa and Navarra. This type of forest is characterized by the dominance of the
holm oak Quercus ilex L. subsp. Ilex. At the time of the research, the stands were
in a recovery process due to the abandonment of charcoal extraction and their development on lands of little economic interest. Although the spider community of
these forests had already been described (CASTRO, 2004b), the seasonal dynamics
of the species had not been analysed, and they were therefore included in this
work. A detailed description of the sampled stands may be found in CASTRO
(2004b).
Igara: This sampling took place in a small eutrophic alder forest (with dominance
of Alnus glutinosa (L.) Gaertner) located outside the city of San Sebastián. The forest
was unexploited, but at a young stage, with no tree measuring more than 35 cm in
diameter at breast height. The understorey was very dense, with several trees covered by ivy (Hedera helix L.) and the presence of scattered fallen trees. The main aim
of this sampling was to compare two different bark trap designs to study trunk-dwelling spiders. The results are currently being analysed and are to be published
shortly.
MATERIALS
AND METHODS
Sampling methods: Table II summarizes the different sampling programs conducted at each forest site. Spider catches come from eight different collecting methods:
86
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
- Kempson method: On each sampling date, 314 cm2 samples of the litter layer,
including the first centimetre of mineral soil, were taken at each forest stand. A steel
cylinder (20 cm in diameter and 25 cm high), equipped with an opening device,
enabled extraction of intact sample units.
Spiders were extracted by creating gradients of light, humidity and temperature
using a Kempson apparatus (KEMPSON et al., 1963). Kempson extraction estimates
densities of spiders and soil fauna with a high degree of accuracy (EDWARDS &
FLETCHER, 1971). A detailed description of the use of this method may be found
in BARRAQUETA (1985).
- Berlese Funnels: Litter samples of 500 cm3, including the first centimetre of mineral soil, were taken with a shovel. Saxycolous moss samples of the same volume
were collected by hand. A plastic recipient was kept in contact with the stone wall
and just under the moss sampled to prevent the individuals escaping. Since there
were no stones above ground level in the Zumaia stand, 8 samples of litter were
taken per sampling date (see Table II).
Each sample was put into a plastic funnel with a 6 mm wire mesh inside and
covered by a fibreglass mosquito net. A 60 W bulb was placed 10 cm above each
funnel and left on for 8 days. This length of time ensures effective extraction (DUFFEY, 1972; HUHTA, 1972). The specimens were collected in glass bottles filled with
a solution of 70% alcohol plus some drops of glycerine inside as a preservative.
More details are given in CASTRO (2004b).
This kind of extraction is one of the most effective for catching spiders
(EDWARDS & FLETCHER, 1971). Berlese funnels are useful for estimating population densities and for studying microhabitat preferences (HUHTA, 1971; DUFFEY,
1972; CANARD, 1981; HÖVEMEYER & STIPPICH, 2000).
- Epigeal pitfall traps: These were made using plastic beakers 6.5 cm in diameter
and 8 cm high. Another smaller beaker was placed inside to avoid turning over mud
and litter around the trap. The risk of flooding by rain and clogging with litterfall was
prevented by making two small holes in the upper third of the beaker, and placing
a cork (evergreen oak forests) or plastic (Artikutza) roof, held by wire, 5 cm above
the trap. Each trap was filled to one-third full with a solution of 4% formalin as preservative. Some drops of detergent were added to increase capture efficiency (TOPPING & LUFF, 1995). Samples were taken uninterruptedly every two weeks, except
in 1996 (Artikutza) on the following dates: 18-II, 28-VII, 8-IX, 3-XI and 29-XII.
Pitfall traps are efficient for collecting a great number of species (UETZ & UNZICKER, 1976; CANARD, 1981; CHURCHILL & ARTHUR, 1999; HÖVEMEYER & STIPPICH, 2000, STANDEN, 2000; BUDDLE & HAMMOND, 2003). They mostly catch
mature individuals (TOPPING & SUNDERLAND, 1992), which facilitates identification at species level of most specimens. They are also useful for studying the seasonal dynamics and mating periods of epigeal spiders, favouring the capture of cur-
87
A. CASTRO GIL
sorial species (CURTIS, 1980; BARRIENTOS, 1985a; CHURCHILL, 1993; LANG, 2000).
Another advantage is that they act continuously, allowing both diurnal and nocturnal species to be caught.
- Malaise Traps: The model proposed by TOWNES (1972) was used: black with a
white roof and thin mesh. The collecting recipient was placed at a height of 2 m
and filled with a solution of 75% ethanol and 5% acetic acid (MARTÍNEZ DE
MURGUÍA, 2002). Samples were taken uninterruptedly every two weeks, except in
1996 on the following dates: 18-II, 28-VII, 8-IX, 3-XI and 29-XII.
These traps are normally used to capture flying insects (PUJADE, 1996). Because
they are complex to install, and collect a considerable quantity of insects, they are
usually set up in small numbers or for short time periods, yielding low catches of
spiders (see data in BARRIENTOS & PUJADE, 1999 and VECÍN et al., 2002). However,
6 traps placed in Artikutza that were active for two years yielded a large number of
spiders. The method biases towards typical dwellers of higher-than-epigeal vegetation strata (HAUGE & MIDTGAARD, 1986). The spider fauna of Malaise traps shows
greater similarity with arboreal stratum than epigeal, because the method probably
collects herbaceous and bush strata dwellers (JENNINGS & HILBURN, 1988).
- Bark traps: These traps were 20 x 30 cm in size. They were tied with wire around
tree trunks at a height of 1.5 m. Two kinds of traps were laid: some made of corrugated cardboard (described in DUFFEY, 1969) and some of grooved plastic (see
CASTRO, 2004b). In the evergreen oak forests, only plastic traps were used, while
in Igara 20 of each type were laid.
Bark traps have been used to study species composition and seasonal dynamics
of spiders on trunks (DUFFEY, 1969; CURTIS & MORTON, 1974; HORVÁTH & SZINETÁR, 1998; HORTON et al., 2001).
- Trunk pitfall traps: These consisted of white plastic beakers 5.5 cm in diameter
and 6.5 cm high. They were tied with wire around tree trunks at a height of 1.5 m.
Two small holes were made in the upper third of each beaker to prevent flooding.
A saturate ClNa solution was used as a preservative. Some drops of detergent were
added to increase catch effectiveness (TOPPING & LUFF, 1995).
These traps are used for the same purposes as epigeal pitfall traps (RUZICKA et
al., 1991; WEISS, 1995; RUZICKA, 1997).
- Beating: This method consisted of beating one sector of vegetation 30 times with
a stick. Spiders were collected on a white sheet of 1 m2 spread out and placed 50
cm under sampled vegetation. Samples were taken between 0.5-1.5 m for liana stratum (made up Smilax aspera L.) and between 1.5-4 m for tree foliage (more details
in CASTRO, 2004b).
Beating allows vegetation parts to be reached that are not accessible using other
methods, yielding an abundant catch. However, it is important to bear in mind that
88
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
the time of day, meteorological conditions and vegetation height can influence the
number of individuals and species composition collected (ABRAHAM, 1983; CODDINGTON et al., 1996; SORENSEN et al., 2002).
- Chance searching by hand: While sampling, spiders detected visually by chance
were collected by hand or with the help of a paintbrush moistened with alcohol.
This method allows valuable complementary information to be gleaned on the
microhabitat and biological aspects of the species (DUFFEY, 1972; FUJII, 1998).
Site
Method
Gorbea
K
Artikutza
P
M
Evergreen
oak forests
P
Fl
Fm
BT
TP
BL
Bf
C
Igara
BT
Number
Number
of
of traps Periodicity collections
or samples
per trap
or sample
Whole sampling
period
Forest
habitats
32-40
1 month
14
V-1982 to VI-1983
Forest litter
30
6
2 weeks
2 weeks
46
46
V-1995 to V-1997
V-1995 to V-1997
Epigeic
-
30
28
20
30
30
30
30
-
2
8
8
4
2
8
8
2
weeks
weeks
weeks
weeks
weeks
weeks
weeks
weeks
27
6
6
12
23
7
7
-
XII-1998 to XII-1999
I-1999 to X-1999
I-1999 to X-1999
I-1999 to XII-1999
I-1999 to XII-1999
XII-1998 to XI-1999
XII-1998 to XI-1999
XII-1998 to XII-1999
Epigeic
Forest litter
Saxycolous moss
Tree trunks
Tree trunks
Liana layer
Tree foliage
Several
40
1 month
12
IX-2002 to VIII-2003
Tree trunks
Table II.- Summary of the sampling program carried out in each study site. Abbreviations: K:
Kempson method, P: epigaeic pitfall traps, M: Malaise traps, Fl: Berlese funnels, litter samples, Fm:
Berlese funnels, saxycolous moss samples, BT: bark traps, TP: trunk pitfall traps, Bl: beating of liana
layer, Bf: beating of tree-foliage, C: chance hand collecting.
Identification and preservation of spiders: Species were identified using the
classic European keys of ROBERTS (1985-87, 1995), HEIMER & NENTWIG (1991),
NENTWIG et al. (2003) and SIMON (1914-1937). The nomenclature used was as
proposed by PLATNICK (2008).
Several more specialized works were used to identify correctly certain individuals
belonging to the following families: Agelenidae: BARRIENTOS (1985b).
Anyphaenidae: URONES et al. (1995b). Atypidae: KRAUS & BAUR (1974) and
CANARD (1983). Linyphiidae: FAGE (1919), SCHENKEL (1938), DENIS (1965), SAARISTO (1974), MILLIDGE (1975), RIBERA & HORMIGA (1985) and BOSMANS
(1995). Mimetidae: CANARD (1982). Salticidae: SNAZELL et al. (1999). Theridiidae:
VANUYTVEN (1991) and KNOFLACH (1993). Thomisidae: LOGUNOV (1992).
Uloboridae: WIEHLE (1964). Zodariidae: BOSMANS (1997).
89
A. CASTRO GIL
The collaboration of several taxonomists was needed to identify some more difficult specimens: Robert Bosmans from the University of Gent, Belgium (Theridiidae
y Salticidae); Miguel Ángel Ferrández from the Society for the Study and
Conservation of Spiders, Spain (Segestriidae, Dysderidae and several families); Peter
Van Helsdingen (Linyphiidae) of the National Museum of Natural History of
Holland, Eduardo Morano of the Iberian Arachnology Group, Spain (Araneidae,
Tetragnathidae); Carles Ribera of the University of Barcelona, Spain (Linyphiidae);
Michael Saaristo of the Museum of Zoology, Turku, Finland (Linyphiidae) and
Carmen Urones of the University of Salamanca, Spain (Miturgidae).
Spiders were preserved in 70% ethanol and stored in the Entomology Department
of the Sociedad de Ciencias Aranzadi (San Sebastián, Spain).
Data analysis: In order to determine which is the best period of the year to sample, seasonal variations in species richness were displayed in graph form. This
determination was based in the highest occurrence of species in adult stage, although species at immature phases were also added to the analysis because there is
little literature available on seasonal variations in their richness. For each species,
graphical displays were limited to the most abundant species (> 10% of identified
sample to the species level per study site and method).
This analysis was limited to sampling methods that were carried out systematically
at least 12 times, completing an entire year. Thus data from samples from Berlese
funnels, beating, and hand searching from evergreen oak forests were used as complementary data to confirm the occurrence of the species in specific periods of the
year and forest strata. Data from the six evergreen oak forests were pooled, and
there are therefore two values for the dates displayed, because in two stands (Ataun
and Larraun) sampling started one week later than in the other four.
The occurrence periods of each species were determined by reviewing data from
this study and from the main studies (101 in total) on biological cycles and seasonal dynamics of European spiders. Among them, several with scattered records were
consulted to complete the scarce knowledge that exists on some little-studied spiders of South European distribution.
Most of the literature consulted was based on pitfall traps. It was therefore possible to determine the main activity periods for several species. For each study consulted, all the maximum activity periods of the most abundant species were registered. Since some species were represented in more than one study and activity patterns may vary from year to year, the period finally obtained for each species is probably wider it would be for any one specific year. Data which did not match the
literature reviewed is dealt with in the discussion.
Classifications of dynamic cycles based on overwinter stages and season and
length of breeding period (TRETZEL, 1954; SCHAEFER, 1977; YSNEL & CANARD,
1990) were avoided for the following reasons: 1) It was impossible to identify all
90
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
immature specimens, 2) there is a lack of information in most literature on immature stages, 3) sometimes the same species is classified differently depending on the
source, and 4) in the absence of direct observations, it is difficult to determine whether a long period of male activity is due to a eurychronous species with several
overlapping generations or to a diplochronous species with a partial concurrence of
two different generations.
RESULTS
Global results and faunistic contribution
Breakdown data of samples from Gorbea, Artikutza and Igara are shown in
Appendices I, II, III and IV, respectively. Breakdown results from evergreen oak forests
have been published in CASTRO (2004b), and for this reason are not given here.
All samples together added up to 6976 specimens collected (inmatures included).
5407 spiders (77.51%) could be identified to species level. The various sampling
methods accounted for the following percentages of identified individuals: Pitfall
traps: 98.44% in Artikutza and 93.74% in evergreen oak forests; Kempson method:
45.51%; Berlese funnels: 82.29%; Trunk pitfall traps: 71.49%; Bark traps: 90.88% in
evergreen oak forests and 91.76% in Igara; Beating: 70.61%; Hand searching:
84.38%.
Table III shows how the higher the forest stratum sampled, the smaller the proportion of adult specimens present, both in abundance and activity. Rates of adults
and immature specimens matched for different sites and years in methods that register abundances in the same kind of microhabitat (Litter: Kempson and Berlese. Tree
trunks: bark traps).
Sampling method (Site)
Males
Females
Juveniles
73.68
52.95
37.40
18.88
15.38
22.79
14.48
15.26
10.94
24.25
48.12
65.86
6.18
7.41
4.06
1.66
2.69
2.75
2.35
15.75
13.33
10.43
13.81
12.94
6.42
7.40
78.07
79.26
85.51
84.53
84.37
90.83
90.25
Activity
P (Artikutza)
P (Evergreen oak forests)
M (Artikutza)
TP (Evergreen oak forests)
Abundance
K (Gorbea)
Fl (Evergreen oak forests)
Fm (Evergreen oak forests)
BT (Evergreen oak forests)
BT (Igara)
Bl (Evergreen oak forests)
Bf (Evergreen oak forests)
Table III.- Percentage of mature and immature specimens caught using each sampling method.
Abbreviations: see Table II.
91
A. CASTRO GIL
Identified specimens have yielded 148 species distributed in 27 families (Table
IV). Three species, Centromerita bicolor, Micrargus apertus and Midia midas were
new records for Iberian fauna, while Peponocranium ludicrum was a new genus
record for Spain. According to PLATNICK (2008), known geographic distributions
for these four species are Palaearctic and Canada, Palaearctic, Europe, and Europe
and Russia, respectively.
Two species Centromerus sp. and Tenuiphantes cf. jacksoni, are currently being
reviewed by specialists. Tenuiphantes cf. jacksoni could be a new species for science (VAN HELSDINGEN, BOSMANS, personal communication).
Following MELIC (2001) – excepting where other authors are indicated – 8 species were endemisms of Ibero-Pyrenean scope or surrounding distributions:
Troglohyphantes furcifer is found in the Spanish autonomous regions of the Basque
Country, La Rioja and Navarra and in the French department of Basses Pyrénées.
Bordea negrei is a Pyrenean endemism. Labulla flahaulti is found in Southern
France and the Pyrenees (HORMIGA & SCHARFF, 2005). Tegenaria inermis,
Chorizomma subterraneum, Walckenaeria dalmasi, and Malthonica lusitanica are
Pyrenean and Northern Iberian Peninsula endemisms, the distribution of the latter
extending to almost the entire Atlantic area of the peninsula (BARRIENTOS & CARDOSO, 2007), while Lepthyphantes bacelarae is an Iberian-Atlantic endemism (CASTRO & ALBERDI, 2002).
Another six species showed a known South-western European distribution:
Dysdera fuscipes from the Atlantic seacoast of Northern Portugal and Spain and the
southern half of France (FERRÁNDEZ, 1985b; LE PERU, 2007). Palliduphantes cernuus from the Pyrenees and Corsica (RIBERA & HORMIGA, 1985). Episinus theridioides from the Pyrenees, Corsica and Sardinia (BOSMANS & CASTRO, 2002).
Anyphaena numida from the French department of Pyrénées Orientales, the northern half of the Iberian Peninsula and Algeria (URONES, 1996). Nemesia simoni
from Portugal, Spain and Southern France, and Entelecara aestiva from Spain,
France, Corsica and Italy (LE PERU, 2007; PLATNICK, 2008).
Seasonal dynamic of species richness: Among most species, activity was concentrated from spring to autumn (Figures 1 and 2). A winter minimum was usually
found in January-February. Activity peaked in spring (normally in May-June) and
autumn (varying between September and November). Some relative minima and
maxima were observed between these two peaks. Autumn maxima showed more
species at immature stages than spring peaks. The largest proportion of immature
spider species was found from late summer, continuing right through the autumn.
Spring peaks were almost exclusively made up of species at adult stage. Peaks in
Artikutza obtained in pitfall traps were smaller in the second year than in the first
year (Figure 1). This trend was not observed in samples from Malaise traps.
Epigeal pitfall and Malaise traps showed continuous activity of adults all year
round, except for some sporadic interruptions in winter in Artikutza. In contrast,
activity halted all winter in trunks of trees in evergreen oak forests.
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Figure 1.- Seasonal changes in species richness recorded in the beech forest of Artikutza.
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Figure 2.- Seasonal changes in species richness recorded in Cantabrian evergreen oak forests.
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Figure 3.- Seasonal changes in species richness recorded in the litter layer and in barks.
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Nonetheless, adult-stage species were present all year round in the litter and
tree bark (Figure 3). In these samples, the largest proportion of species in adult
stage also occurred in spring (and in a part of the summer in the evergreen oak
forests). The minima occurred in winter in the bark traps and in summer in
Kempson samples. Overall species richness in bark traps attained a maximum in
autumn, continuing in winter, but interrupted by a relative minimum in January or
in February. In litter samples from Gorbea, overall species richness peaked in
January, showing another relative maximum in April, after a relative minimum in
February-March.
If we continue to focus on adult stages, Table IV shows periods of maximum
activity and presence for each species. It has been possible to determine the
period of maximum activity of males, females, or both for 119 species. May and
June were the months most frequently included in the periods of maximum breeding activity (Figure 4). Together these months encompassed 78.15% of species.
If the entire period of adult presence – currently known – is taken in account,
besides breeding peaks, 93.92% of the 148 species found were included between
May and June.
Sampling data coincided with this trend: May and June usually continue to be
the months with the highest number of species (Table V). Kempson samples from
Gorbea and bark traps from the evergreen oak forests were the exceptions to this
pattern. However, pooled together, May and June ranked as the second richest
Figure 4. Percentage of mature species that include the specified month in their main reproduction
period.
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Table IV.- Months of the highest activity and presence (in brackets) of each species.
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Table IV.- Continue
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Table IV.- Continue
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Table IV.- Continue
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SEASONAL
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Table IV.- Continue
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Table IV.- Continue
period in Gorbea (13 species), and June-July in the oak forests (6 species).
Besides, the lower the stratum, the sooner the species richness peak occurred:
mainly in May in epigeal pitfall traps, June in Malaise and trunk pitfall traps, and
the second half of June and beginning of July in trees (trunk pitfall and bark traps
pooled).
Seasonal dynamics of the species: Seasonal dynamics of the most abundant
species are displayed as graphs in Figures 5-9. Species whose seasonal dynamics are
poorly known or for which scarce literature data are available, are shown in Figures
5 and 6. Better known species are represented in figures 7-9.
For some species, males were active for a long period (Figure 5): Adults of
Malthonica lusitanica show oscillations in activity, though this occurred mainly in
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SEASONAL
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spring and autumn, with a peak of males in the autumn. Palliduphantes cernuus
was active all year, except for late fall and the beginning of the winter, reaching the
maximum in spring and early summer. Adults and immature specimens of
Chorizomma subterraneum were present all year round. Adults were active mainly
in autumn and winter, though with several fluctuations. Juveniles are most active
from mid-summer to the beginning of the autumn, with three peaks of abundance
in late spring, the first half of the autumn and winter (the maximum). Maximum
adult abundance of Gongylidiellum murcidum occurred in late autumn and the
beginning of the winter. Although males could be found for all autumn and winter,
this period did not perhaps indicate true mating activity (this issue is dealt with later
in the discussion). Other species with long activity periods in males (see Figure 7
and Appendices II and III) were Tenuiphantes flavipes (spring to autumn),
Tenuiphantes zimmermanni (spring and summer) and Tenuiphantes cf. jacksoni
(spring and summer).
Other species showed male activity periods restricted to a certain time of year
(Figure 6): In Dipoena melanogaster this was from late spring to the beginning of
summer. Labulla flahaulti adults were present from the second half of the summer
to mid-autumn, while immatures appear in spring and summer. Nemesia simoni
males reached maximum activity in autumn, mainly between September and
November, with a few sporadic adult catches during the rest of the year. Males of
better-known species (Figure 7 and Appendices II-IV) were active in the following
periods: Clubiona comta, Anyphaena accentuata and Pardosa lugubris in spring,
Malthonica picta in spring and the beginning of summer, Textrix denticulata and
Clubiona terrestris in late spring and summer, Coelotes terrestris from mid-summer
to mid-autumn and Scotina celans in late autumn and winter.
There were species in which both adult males and females, were present in two
different periods of the year: Episinus maculipes at the beginning of spring and in
summer (Figure 5), and Micrargus apertus (Figure 6) in spring and late autumn-winter.
Bark traps showed the seasonal dynamics of immatures of several species, whose
captures were concentrated from late summer to winter (Figure 8). Despite the activity overlap among the species, their peaks did not coincide, with the exception
of Clubiona brevipes and Cheiracanthium mildei. However, both species co-occurred in the forest of Zumaia, where the dates of their peaks also differ (CASTRO,
2004b).
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Figure 5.- Seasonal dynamics of species with scarce bibliographical data.
SEASONAL
DYNAMICS OF FOREST SPIDERS
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BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Figure 6.- Seasonal dynamics of species with scarce bibliographical data.
A. CASTRO GIL
Figure 7.- Seasonal dynamics of species collected in high numbers in Malaise traps and epigeal
pitfall traps (Artikutza).
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DYNAMICS OF FOREST SPIDERS
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Figure 8.- Seasonal dynamics of immature specimens of species caught in high numbers using bark
traps.
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Figure 9.- Seasonal dynamics of some species during two consecutive sampling years (Artikutza:
Malaise traps).
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SEASONAL
DYNAMICS OF FOREST SPIDERS
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Vertical distribution of the species: Several species were active in more than
one forest stratum. This was the case of Malthonica picta, Tenuiphantes cf. jacksoni, T. zimmermanni (Figure 7) and T. flavipes. In Artikutza, these species were
caught both in pitfall and Malaise traps. But in the evergreen oak forests, they were
not collected, or only sporadically collected, in trunk pitfall traps and bark traps,
and never reached the tree foliage and liana stratum. In contrast, Clubiona comta
and Saitis barbipes were caught by all kinds of collecting methods. Dysdera fuscipes, Harpactea hombergi, Neon robustus and Saitis barbipes were also active in both
Site and Whole
sampling sampling
period
methods
Dates that showed the
highest species richness
The two consecutive dates that
showed the highest species richness
Richness
Dates
Richness
%
Dates
Richness
%
25
I
11
44.00
XII-I
14
56.00
30
45
62
15 V
26 VI
15 V
18
14
25
60.00
31.11
40.32
15-29 V
26 VI-10 VII
15 V
22
21
35
73.33
46.67
56.45
16
42
47
20 IV
9 VI
9 VI
7
16
17
43.75
38.10
36.17
6-20 IV
26 V-9 VI
26 V-9 VI
7
21
22
43.75
50.00
46.81
32
61
74
15/12 V
12/9 VI
29/26 V
18
20
28
56.25
32.79
37.84
15/12-29/26 V
29/26 V-12/9 VI
29/26 V-12/9 VI
22
30
41
68.75
49.18
55.41
P
TP
BT
54
30
17
22/29 V
5/12 VI
28 VIII/4 XI
and 25 IX/2 X
16
9
5
29.63
30.00
29.41
22
13
7
40.74
43.33
41.18
P + TP
P + BT
73
63
5/12 VI
3/10 VII
22
18
30.14
8.57
30
22
41.10
34.92
TP + BT
P + TP + BT
39
80
3/10 VII
3/10 VII
10
24
25.64
30.00
8/15-22/29 V
5/12-19/26 VI
3/10 VII-28
VIII/4IX and
28 VIII/4 IX25 IX/2 X
5/12-19/26 VI
8/15-22/29 V and
22/29 V-5/12 VI
3/10-17/24 VII
5/12-19/26 VI and
19/26 VI-3/10 VII
14
30
35.90
37.50
12
XI and V
4
33.00
V-VI
6
50.00
Gorbea
K
Artikutza
First cycle
P
M
P+M
Second cycle
P
M
P+M
Total
P
M
P+M
Evergreen
oak forests
Igara
BT
Table V.- Observed species richness (only in adult phase) with each sampling method (or by combining more than one) in the whole sampling period and during the periods of highest species richness. Abbreviations: see Table II. In Artikutza, the first cycle ran from 15 May 1995 to 28 April 1996,
and the second from 12 May 1996 to 20 April 1997.
109
A. CASTRO GIL
environments, epigeal and tree trunks and the latter extended its activity to tree
foliage. Despite showing vertical displacement, other species such as
Palliduphantes cernuus, showed greater activity in lower strata, while Episinus
maculipes, Labulla flahaulti and Textrix denticulata were mainly found in upper
strata.
All the most abundant species with long mating period (2 seasons or more) were
found in the coger vegetation strata. The only exception was Tenuiphantes zimmermanni. However, this species was only caught in large numbers in Malaise
traps, being sporadically found in bark traps and totally absent in trunk pitfall traps
and beating samples.
Differences between consecutive years: In Artikutza, there were differences
in abundance within species between the first and the second year (Appendices II
and III). Higher numbers of individuals were registered by pitfall traps in the first
year for almost all the most abundant species: Malthonica picta, M. lusitanica,
Coelotes terrestris, Tenuiphantes cf. jacksoni, T. zimmermanni and Pardosa lugubris. The only exception was Nemesia simoni, where the trend was just the opposite (Figure 6). On the contrary, with the exception of Tenuiphantes zimmermanni, catches in Malaise traps were more numerous on the second year for the most
abundant species: Malthonica picta, Textrix denticulata, Clubiona comta and
Clubiona terrestris.
Within each species, main male activity did not vary more than one month between different years and places. In Nemesia simoni (Figure 6), the main activity
always took place in October (though it extended to November in 1996). In Coelotes
terrestris this period occurred in September; in Malthonica picta (Figure 7) and
Pardosa lugubris it varied from April to May; and in Clubiona terrestris and Textrix
denticulata (Figure 9), it took place from July to August (but it is possible that the
July peak did not vary, given that in the second year, a July sample was missing in
Artikutza). Only Clubiona comta showed larger oscillations, with the main male
activity between late March and mid-May depending on the year. However, a lack
of samples before May in 1995 in Artikutza may have led to an earlier activity peak
for this species being missed.
DISCUSSION
Faunistic contribution: The capture of the following new species for the
Iberian Peninsula and Spain was not surprising, since all are widespread in Europe
(CANARD, 2005), and have already been recorded from the south of France:
Centromerita bicolor had been registered from several locations in the French
Pyrenees (BOSMANS & DE KEER, 1985; LEDOUX et al., 1996; LE PERU, 2007),
Peponocranium ludicrum is common in the Atlantic side of France (LE PERU, 2007)
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and has been recently recorded in Portugal (CARDOSO et al., 2008). Micrargus
apertus is a less frequent species, usually recorded in forest ecosystems in Europe
(HÄNGGI et al., 1995), but in France it is only known in the Department of Pyrénées
Orientales (MILLIDGE, 1975).
The nearest record for the fourth species, Midia midas, is in a forest in the
Department of Seine-et-Marne (SIMON, 1884, 1914-1937), in the north of France.
There are more scattered citations of Midia midas in Central and Atlantic Europe
(CANARD, 2005). The species is considered an old growth forest specialist (DUFFEY, 1993; HARVEY et al., 2003). According to these authors, Midia midas seems to
be confined to senescent trees, living in loose bark, dead wood, litter inside cavities of beeches and oaks, and in the nests of birds and squirrels. Corroborating these
data, Midia midas was found in the forest of Artikutza, the only one with a large
amount of dead wood caused by a lack of forestry work over more than 70 years
at the time of sampling (MARTÍNEZ DE MURGUÍA et al., 2002). Only one specimen
was found, collected by Malaise traps. More specimens would probably have been
found if the oldest trees in the forest had been sampled.
Despite finding 14 endemic species or with South-western European distribution,
their presence in the study area was expected, because it is included in their known
geographic distribution.In any case, the capture of Bordea negrei is remarkable,
since hitherto all known citations have been from caves (BOSMANS, 1995).
Surprisingly, the only individual caught was found in a pitfall trap in Artikutza,
where the soil has no connection to a subterranean cavernicolous environment,
because of the substrate of intrusive rocks characteristic of this area (CATALÁN et
al., 1989).
Seasonal dynamics: In agreement to the literature reviewed the species richness peak of active adult specimens takes place in the temperate part of the central
and atlantic Europe between May and June: In Germany in several strata (TRETZEL,
1954), in Denmark for several strata pooled (TOFT, 1976), and in Portugal in epigeal stratum (CARDOSO, 2004; CARDOSO et al., 2007). However, this pattern is broken in mediterranean places as in west-central Spain, where the main epigeal activity takes place in June-August, with the maxima in August (JERARDINO et al.,
1988). Likewise, in boreal climates like in Finland, there is also a delay in the peak
of species richness, which occurs between June and July (NIEMELÄ, 1994).
The later occurrence of maximum richness in upper strata has also previously
been observed: In German forests, epigaeic richness extends from April to June, and
trunk-canopy richness from June to July (ALBERT, 1982). LUCZAK (1959), sampling
the ground flora with a sweep net in Polish pine forests, found June to be the period
of main diversity. HOREGOTT (1960) sampled the arboreal stratum of a German
pine forest and concluded that the diversity peaks in July-August. Although they do
not provide data on seasonal variations in species richness, other studies report the
111
A. CASTRO GIL
same months within the peaks of maximum adult spider abundance: in Germany,
May-September in the epigeal stratum and May-July in the trees (ALBERT, 1982) and
in Austria May-July in the epigeal stratum (NOFLATSCHER, 1993).
All of these results were expected, since most species breed between the spring
and summer in most European countries: Germany (SCHAEFER, 1977), Austria
(NOFLATSCHER, 1993), Belgium (BAERT & KEKENBOSCH, 1982; BAERT et al.,
1983), Finland (PALMGREN, 1972) and Norway (HAUGE, 2000).
In this work, whereas the May-June peak is due almost exclusively to the activity
of adult spiders, the second relative maximum, usually in autumn, is the result of
active immatures in period of dispersal, most probably looking for overwintering
shelters (SCHAEFER, 1977). These immatures probably come from females that
mated in spring and summer (LUCZAK, 1959). At this time, there are also some
adults in breeding activity and others looking for sites in which to overwinter
(TOFT, 1976; SCHAEFER, 1987). The sum total of the adults’ activity causes the
autumn peak observed in Artikutza. This second maximum has also been observed
in several localities in Portugal, specifically in October (CARDOSO, 2004; CARDOSO et al., 2007).
Results from litter samples and bark traps reflect other dynamics apart from seasonal reproductive displacements. For example, linyphiids are the most abundant
and diverse family in litter samples in Gorbea. Among them, several species breed
in the fall or winter (SCHAEFER, 1976; CARDOSO, 2004). In addition, other species
overwinter in the litter (SCHAEFER, 1977), explaining the winter maximum observed.
The summer minimum coincides with the driest season. This condition reduces the
presence of linyphiids (HAUGE, 2000). Summer is also the season when the least
amount of litter is available on the forest floor (GABBUT, 1956) and higher amounts
of litter are usually positively linked with diversity in of spiders (UETZ, 1979).
The autumn movements of species looking for overwintering sites (DUFFEY,
1969; PEKÁR, 1999b; HORTON et al., 2001; HORVÁTH & SZINETÁR, 2002) are responsible for the maximum species richness observed in this season using bark traps.
As the each species’ activity ends, diversity falls during the winter, to a JanuaryFebruary minimum. From then on, there is activity among more species in adult stages, and thus juveniles mature or disperse to other strata or environments. As a
result, there was a second peak of adults in spring-summer. During the warm season some species breeded or feeded in tree trunks: Textrix denticulata, Harpactea
hombergi, Clubiona comta, Labulla flahaulti, Segestria bavarica and Neon robustus.
Some of them were not present in the Igara samples, causing a relative summer
minimum. In the autumn, several of these species disappeared and displacements
of immatures looking for overwintering sites started again.
Seasonal dynamics of the species: Usually, species with long-reproductive
periods do not concentrate them in one specific time of the year (SCHAEFER, 1987).
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This is the example of the mating period of Tenuiphantes flavipes, as observed by
several authors (MERRETT, 1969; JOCQUÉ, 1973; RIBERA & HORMIGA, 1985). The
species appears to be very flexible in this regard since peaks of males can occur at
different times of year, according to different studies: winter (TOFT, 1976), June
(SCHAEFER, 1976), and even in two different periods of the year, May-July and
October-November (HARVEY et al., 2002). The principal mating period of
Palliduphantes cernuus extends from late spring to the beginning of summer, according to specimens collected in the Pyrenean foothills, Jaca, Spain (RIBERA & HORMIGA, 1985). But in Jaca, the species shows two differences with regard to this
study: a second peak of males between December-February, and an adult minimum
extending from August to November. There are also differences between the two
Spanish areas in the seasonal dynamics of Chorizomma subterraneum. In Jaca, the
mating period is between August and December, with a maximum between August
and October (BARRIENTOS, 1985b). But in the study area this period extends to
May, with three maxima observed: September-October, December-January (peak of
male activity), and February-April (secondary peak interrupted by a slight drop in
March). In Jaca, Chorizomma subterraneum has been collected in damp low-altitude forests, and is more common in floors with thick layers of moss and needles.
This, combined with its troglophilous character (RIBERA, 1980), leads one to think
that Chorizomma subterraneum is a stenotopic species looking for damp habitats
with narrow temperature margins throughout the year (BARRIENTOS, 1985b). The
colder winter of Jaca would probably prevent activity of the species, at least on the
epigaeic layer. In the Basque Country, its life cycle seems to have an overlap of
three generations per year, thus each mating peak is followed by an activity of females and a consecutive peak in abundance of immatures, which remain inactive in
the litter during the cold season (Figure 5).
Data from the literature shows greater consensus in the case of Tenuiphantes zimmermanni and Malthonica lusitanica. Both species have a minimum reproductive
activity in winter, with two main mating periods between spring-summer and late
summer-autumn. However, in Malthonica lusitanica, the spring-summer activity of
males can be greater—as in Portugal (BARRIENTOS & CARDOSO, 2007) and
Artikutza—or smaller than the late summer-autumn peak, as in Jaca, Spain
(BARRIENTOS, 1985a,b) and the evergreen oak forests. In addition, BARRIENTOS
(1985b) observed differences between habitats sampled at high and low altitudes
within the same geographic area: at higher altitudes, males did not show springsummer activity. Hence, what is always conservative is the late summer-autumn
peak. BARRIENTOS (1985b) suggests that winter and spring breeding activity might
be confined to deeper layers inside the litter. It has been observed that Malthonica
lusitanica shows a clear preference for woodland habitats with a relatively deep litter and ground moss (BARRIENTOS & CARDOSO, 2007). In the litter and saxycolous moss of evergreen oak forests, adults are mainly concentrated in spring-sum-
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A. CASTRO GIL
mer (males only in spring), and immatures all year round (CASTRO, 2004b). In this
kind of forest, pitfall traps show that each peak in males, even secondary ones, is
followed by activity amongst females. These data suggest the presence of consecutive overlapping generations, as previously hypothesized (BARRIENTOS, 1985b).
Among species with short reproductive periods, only Scotina celans showed a
peak in male activity that did not coincide with the data from the literature.
According to the references consulted, the peak occurred between September and
November, while in Cantabrian evergreen oak forests it occurs between December
and February, with a presence stretching to April. Females were found all year
round. The same was true for immatures, which abundantly colonized the saxycolous moss. In Brittany (France), CANARD (1984) found that adults started to be active at the beginning of the autumn, and the greatest population of this species had
a biological cycle of two years, it being possible that one small portion completed
it in a single year. It is possible that in the more moderate climate of the evergreen
oak forests of the study area, the cycle could be mostly resumed in one year, with
the first males maturing in the second half of the autumn. Only URONES (1985a)
also finds a persistence of males from October to April in the Pyrenean foothills
(northern Spain), but her data is made up of a small number of specimens, and they
come from different habitats comprising a wide spectra of climatic conditions that
may represent a mix of one and two-year cycles.
Within a given species, peaks of activity and density are often not coincident
(HUHTA, 1965; 1971). This is the reason for the differences usually observed in the
results obtained by pitfall traps and extraction of litter samples (HOVEMEYER &
STIPPICH, 2000). For instance, Gongylidiellum murcidum shows a spring-summer
breeding period, the male peak occurs in summer, and the presence of adults
extends till the autumn (PALMGREN, 1972, 1976; PLATEN et al., 1996). The data in
PLATEN et al. are based on pitfall traps, and PALGREM’s on litter sifting in the field
throughout the year with the exception of the winter. According to the data collected in litter samples in Gorbea (Appendix I), males occurred in autumn and winter,
and females all year round, with a peak of adults in January. This peak is in concordance with that obtained by WOZNY (1992) in the litter and moss of Polish pine
forests. But this author does not break down the data into males and females. A possible interpretation of all these results may therefore be summarized as follows:
there is activity of adults from the spring; the females that mate earlier could lay
eggs from which spiders would hatch that become mature in the autumn. These
individuals would overwinter—mainly inactively—in the litter, and be active again
in spring and summer. This would explain the drop in density observed in Kempson
litter samples, because litter extraction methods bias, above all, spiders that are inactive or show static hunting strategies (KOPONEN, 1976). The seasonal dynamics of
Micrargus apertus were similar (Figure 6). According to HARVEY et al. (2002), in
England the main breeding period of M. apertus is in spring and early summer, with
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sporadic presence of adults all year round. Even though they do not specify the
sampling methods used to collect this species, it is likely that they reflect its activity.
This winter activity, on mild, sunny days, is common in Central Europe (KIRCHNER,
1987). In accordance with the data obtained by BARRIENTOS et al. (1994),
Harpactea hombergi is another species whose adults are mainly inactive in winter,
but it overwinters in the evergreen oak forests, especially in the saxycolous moss.
Bark traps show displacements of spiders searching for overwintering sites (DUFFEY, 1969). As this researcher found, I observed that Clubiona comta also used bark
traps for egg-laying (females with eggs observed in June and July samples), and C.
brevipes as an overwintering site. In addition, Malaise traps shows that immatures
of C. comta have more winter activity, being found in the litter, tree foliage, and
mainly in tree trunks and arbustive stratum. Corroborating the data showed by
KOOMEN (1998), immatures of Anyphaena accentuata were also active in winter,
but more restricted to the arboricolous environment. As the findings of HORTON et
al. (2001) and HORVÁRTH & SZINETÁR (2002) show, juveniles of C. pallidula and
Cheiracanthium mildei also rested in the traps, being sporadically active in winter.
Although most individuals from all these species overwinter as immatures, a few do
so as adults. Presence of C. terrestris adults has been found in November, C. comta
and C. brevipes till December, and A. accentuata in January. However, winter activity of adults has not been observed. The mere presence of adults in winter for
some of these species is documented in the literature: C. comta (SCHAEFER, 1976;
BARRIENTOS et al., 1996; HARVEY et al., 2002), C. pallidula (SCHAEFER, 1976), C.
terrestris (SCHAEFER, 1976; TOFT, 1976, 1978a; HARVEY et al., 2002), and A. accentuata (URONES et al., 1995a; HARVEY et al., 2002). On the other hand, Clubionidae,
Miturgidae and Anyphaenidae spiders are all cursorial spiders. The lack of coincidence among activity peaks of spiders belonging to the same guild has been observed previously, and it supports the hypothesis of temporal stratification as a possible factor that explains their co-existence (TRETZEL, 1954; BUDDLE & DRANEY,
2004). However, at least within Clubionidae there is no evidence of intraguild predation on tree trunks (PEKÁR, 1999b). Hence, future research will be needed to test
this hypothesis in the study area.
Because their life cycles are not well-known, a further discussion of the data
obtained for the species Nemesia simoni, Labulla flahaulti, Dipoena melanogaster
and Episinus maculipes is presented below:
Hitherto, sporadic records have given clues about the mating period of Nemesia
simoni. The anecdotic observations of THOMAS (1999), who found 46 males (and
any females) drowned in a swimming pool in the Department of Gironde (France)
in a period of less than five days (12-16 September 1994), tallies with the data obtained in this work, where males of Nemesia simoni concentrate their activity from
September to November, with a low number of additional captures till January in
the evergreen oak forests.
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The cycle of Labulla flahaulti fits with that of autumn stenochronous (SCHAEFER,
1977). Immatures were collected from late March, showing activity till August
(Figure 6), when more individuals became sub-adults or adults. Throughout that
time, they build their webs between large epigeal stones, barks and foliage of trees
and bushes (CASTRO, 2004b). Adults are mainly found in tree trunks. Mating took
place between the middle of summer and early autumn, when it was common to
observe webs of both, males and females, in the same bark trap (personal observation). An egg-sac, found in Igara samples in October, suggests that autumn is the
egg-laying period. No specimens were collected in winter, which indicates an
annual cycle, overwintering in the egg phase.
Dipoena melanogaster shows a life cycle that falls into the spring and summer stenochronous category (SCHAEFER, 1977), which matches the findings of PLATEN et
al. (1996). The mating period took place between May and July, with a peak in
June. BARRIENTOS et al. (1996) indicate that mating occurs in summer, but without
specifying which months. This species - only found in the Cantabrian evergreen oak
forests in the present work – overwinters as immature stage in the foliage and branches of trees and shrubs, after showing a remarkable period of activity in the fall.
In the Cantabrian evergreen oak forests, Episinus maculipes males occurred in
two different seasons: spring (from April to May) and summer (July-September).
Adult females have also been registered at the same times. But in the
Mediterranean evergreen oak forests of North-Eastern Spain, the mating period
only takes place in summer (BARRIENTOS et al., 1996). The species may show
diplochrony depending on environmental conditions. The immatures were active
in spring and summer on tree trunks, and in the autumn on forest litter where they
probably overwinter. However, no individuals have been collected in litter samples, but several have been observed in branches near the forest floor in the cold
season of the year.
Vertical distribution of the species: The greater proportion of juvenile specimens found in higher strata is explained by the scarcity of species with long breeding periods and by the low winter activity of adult individuals compared with the
epigeal stratum. The higher proportion of species active in the adult stage at the
lowest strata has previously been observed by several researchers (TRETZEL, 1954;
PALMGREN, 1972; TOFT, 1976; ALBERT, 1982; CASTRO, 2004b). PALMGREN (1972)
hypothesised that this pattern is due to the greater thermal stability of lower strata.
HUHTA (1965) and BARRAQUETA (1985) show that air temperature oscillates more
than that of the humus. Likewise, BRAUN (1992) found that higher parts of the trees
are colder and drier, showing less diversity of spiders. MERRETT (1968) and TOFT
(1976) argue that breeding periods are synchronized within each stratum with the
period of maximum activity of prey. According to this, it has been observed that
prey abundance is correlated with species richness of litter spiders, above all in
spring and early summer (UETZ, 1975, 1979). Precisely, the prey capture by litter
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DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
spiders is the greatest in spring months, a period of rapid growth and maturity for
many species (MOULDER & REICHLE, 1972).
We can distinguish between the preferences of several spiders for specific strata.
Like Tenuiphantes flavipes and T. zimmermanni (JOCQUÉ, 1973; TOFT, 1976),
Tenuiphantes cf. jacksoni seems to colonize the plant strata just above the forest
floor surface, as Malaise traps indicate. But none of these species seems to show an
affinity for tree trunks, since they have rarely been collected in bark and trunk pitfall traps. Data from the literature (ALBERT, 1976; WUNDERLICH, 1982; MARC,
1990; BRAUN, 1992; HORVÁTH & SZINETÁR, 1998, 2002) corroborates this hypothesis for T. flavipes and T. zimmermanni. Although T. flavipes can spread to tree
trunks, it prefers the epigeal stratum (SIMON, 1991). Taking pitfall traps as well, it
is observed that Malthonica picta is active in both strata, and that Coelotes terrestris
makes some vertical displacements, even to tree trunks (ALBERT, 1976). Even
though Clubiona terrestris was only well represented in Malaise traps, TOFT (1978a)
finds that it distributes in all strata in a Danish beech forest, as it was the case with
C. comta in this study. Among the most abundant species found, the following are
considered to be at least facultative trunk dwellers, according to the aforementioned literature: Harpactea hombergi, present in the forest floor too, Textrix denticulata, which also occurs in lower strata, and Segestria bavarica, Dipoena melanogaster, Episinus maculipes, Paidiscura pallens, Keijia tincta, Labulla flahaulti,
Anyphaena accentuata, Clubiona brevipes, C. pallidula, Philodromus rufus, P.
aureolus, P. dispar and Diaea dorsata, which are equally found in tree trunks, tree
foliage and arbustive strata.
For the reasons explained in the section on material and methods, of all sampling
methods applied, only Malaise traps are not of widespread use among arachnologists. It therefore seems appropriate to discuss some of the properties of the method. According to the results observed, Malaise traps yield a high number of species, requiring active displacements of spiders to allow their capture. The faunistic
composition recorded seems to correspond to that of the herb and bush forest strata surrounding the traps, as also observed by JENNINGS & HILBURN (1988); it is
not possible to relate the species collected with any particular microhabitat. All these
properties suggest that pitfall and Malaise traps perform similarly and therefore the
same remarks on the interpretation of the results obtained apply to both. It is also
important to point out that spiders account for only 0.11-0.12% of all arthropods
captured in Malaise traps (PUJADE, 1996; SCHNEIDER & DUELLI, 1997), and therefore this method is not recommended if the main focus of the study is the spider
fauna. Nonetheless, when available, data from Malaise traps should not be neglected, since in some forests the highest species richness of spiders has been observed
in low understorey strata (TURNBULL, 1960). Of the 77 species collected in the
beech forest of Artikutza, 34 (44.16%) were registered by pitfall traps and 64
(83.12%) by Malaise traps. These percentages tally quite closely with those found in
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a German beech forest (ALBERT, 1976): 42.35% by pitfall traps and 78.82% by arboreal photoeclectors (85 species collected). In the Cantabrian evergreen oak forest
100 species have been found, 66 of which are present in the epigeal strata and 55
above this level (CASTRO, 2004b). The data obtained therefore highlights the need
to sample above epigeal stratum to get a complete inventory of spider fauna.
Regarding future research, it would be advisable to compare Malaise traps with
other sampling methods well recognised as being effective in registering spiders
active in low forest strata, such as emergency traps (ASCASO & BARRIENTOS, 1986;
HÖVEMEYER & STIPPICH, 2000) and arboreal photoeclectors (ALBERT, 1976).
Differences between consecutive years: The differences found in pitfall trap
samples in Artikutza between two consecutive years, both in terms of the number
of specimens and the species observed, may be explained by several factors: one
could be the weather conditions, since the second year was rainier and colder than
the first (MARTÍNEZ DE MURGUÍA et al., 2001). In Finland, KOPONEN et al. (1974)
found a positive correlation between temperature and spider activity, though
NIEMELÄ et al. (1994) do not, even though the latter observe a negative correlation
with rainfall. MERRETT (1968) also observed a decrease in the activity of lycosid spiders during rainy periods. Data from field experiments in a temperate forest in the
eastern USA also show a reduction in activity for gnaphosids in certain litter layers
(LENSING et al., 2005). Although in Portugal, CARDOSO et al. (2007) obtained a
negative correlation between species richness and rainfall, they did not find temperature to have a significant influence. Although temperature and rainfall might
explain the lower activity observed in pitfall traps in the second year, they do not
explain why these differences are not so evident in Malaise traps. Several species
were caught even more abundantly in the second year by Malaise traps. This effect
might be due to the attraction this kind of trap holds for spiders as a sheltering
microclimate, a focus of concentration of potential prey and their complex architecture (JENNINGS & HILBURN, 1988). Pitfall-trap sampling intensity is another factor that might depress the populations of the most active species (CANARD, 1981).
Within short-mating period species, slight differences in the occurrence of peak
activities of males in consecutive years have also been observed by several authors.
In Germany, TRETZEL (1954) found oscillations of one or two months, arguing that
it is due to variations in temperature. AITCHINSON (1984) observed differences of
two weeks in lycosid spiders in Canada, while BUDDLE & DRANEY (2004) recorded even fewer differences for dominant linyphiids in the same country. RUSSELLSMITH & SWANN (1972) compared seasonal peaks from England and Germany,
obtaining differences that did not exceed of one month. As explained previously,
short-mating period species comprised the majority found in the samples, and also
in temperate latitudes, and the life cycle of most of them (spring and summer stenochronous and diplochronous) is controlled by photoperiod, besides temperature
(SCHAEFER, 1977; KISS & SAMU, 2002). This would explain why the activity peaks
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SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
are so conservative, and why the main predictive factor of species richness over
time is the length of the day, as found by CARDOSO et al. (2007).
Activity minima took place in winter, coinciding with the coldest months of the
year in the study area: January and February (MARTÍNEZ DE MURGUÍA et al., 2001;
CASTRO, 2004b). These months are usually rainy, but the winter of 1997 was warmer and drier than the previous year in Artikutza (MARTÍNEZ DE MURGUÍA et al.,
2001). This would explain the relative maximum observed in the winter of 1997. The
presence of relative maxima in winter in the study area is not surprising, given the
activity of spiders observed in boreal regions on mild days at this time of the year
(HUHTA & VIRAMO, 1979; AITCHINSON, 1984). Species whose activity is more
dependent on temperature, or on complex interactions between temperature and
photoperiod, might interrupt their simple dormancy in warmer-than-usual days in
winter (SCHAEFER, 1977).
Conclusions and further research: With the aim of applying the indicative
value of spider diversity in environmental management and conservation, a thorough knowledge is needed of the representative fauna of the forests in the study
area. To save resources, budget and time, samples should be concentrated in the
periods of the year that yield the most productive results. Most European literature
on the topic suggests that peaks of species-richness at adult stage, and consequently
the highest proportion of identifiable specimens, occur in May-June. In the study
area, in the transition between these months (4 weeks), it is possible to collect
around half of the species recorded in a systematic sampling comprising an entire
year (similar to the percentages found by SCHARFF et al. (2003) and CARDOSO et
al. (2007)), though according to the literature revised, it would theoretically be possible to collect a proportion closer to 94%.
However, as SCHARFF et al. (2003) show, for a specific month, even short intensive three-day sampling protocols can yield results that are as good as two months
of systematic collection carried out at two-week intervals. Thus, for future studies,
the available data from the Cantabrian evergreen oak forests and Artikutza will make
it possible to estimate the performance of short-intensive sampling protocols proposed by several researchers (CODDINGTON et al., 1991, 1996; SORENSEN et al., 2002;
SCHARFF et al., 2003; CARDOSO et al., 2008). If these protocols prove effective, considerable material and economic resources could be saved in future research.
It should also be borne in mind that as observed in Artikutza, both the number
of individuals and the species richness registered may vary between consecutive
years in the same study site. For this reason, this study can be used as a starting
reference work that can be used to check whether the yield for a specific year in
future short intensive sampling protocols is significantly lower than expected.
Finally, another factor to be taken into account is the forest stratum. Results indicate May (the middle half as the most favourable period) as being the best month
119
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to capture the highest species richness in the epigeal stratum (pitfall traps), the MayJune transition for the low understorey vegetation (Malaise traps), and June for tree
trunks (trunk pitfall and bark traps). When limited resources lead to a focus on a
specific stratum, sampling the epigaeic community seems the advisable option in
Mediterranean-type forests in the Iberian Peninsula, since it contains the highest
diversity and rate of endemics (CASTRO, 2004b, CARDOSO et al., 2007).
Nevertheless, this may not be generalized to temperate woodlands since more species richness has been observed in upper strata in Artikutza and other beech forests
(ALBERT, 1976). Further research combining several sampling methods in different
kinds of forests in the study area is needed to determine whether most diversity and
endemic species are generally concentrated in the epigeal stratum.
ACKNOWLEDGEMENTS
This work was assisted by a grant for young researchers conceded by the Society
of Sciences of Aranzadi.
Pilar Barraqueta (EKOS, Environmental Advice and Research of Amorebieta,
Spain), Leticia Martínez de Murguía (Society of Sciences of Aranzadi, San Sebastián,
Spain) and Alazne C. Uribe-Etxeberria (Haritzalde Naturalist Association San
Sebastián, Spain) donated their spiders´collections to carry out this study.
The following students, Ainhoa Iraola, Oneka Zapirain (University of the Basque
Country, Spain) and Jagoba Malumbres (Autonomous University of Barcelona,
Spain), collaborated in the separation and identification of spiders.
Robert Bosmans (University of Gent, Belgium), Miguel Angel Ferrández (Society
for the Spiders´ Research and Conservation, Spain), Peter Van Helsdingen (National
Museum of Natural History, Holland), Eduardo Morano (Iberian Group of
Arachnology, Spain), Carles Ribera (University of Barcelona, Spain), Michael Saaristo
(Zoological Museum of Turku, Finland) and Carmen Urones (University of
Salamanca, Spain) revised and identified some specimens.
Christian Kehlmaier (Museum of Natural History of Dresde, Germany), Javier
Barriga (Complutense University of Madrid, Spain), Alberto Jiménez (National
Museum of Natural History of Madrid, Spain), Antonio Melic (Iberian Group of
Arachnology, Spain), Alberto López Pancorbo and Eva de Más (University of
Barcelona, Spain) helped me with the bibliographical search.
BIBLIOGRAPHY
•
120
1. ABRAHAM, B. J. 1983. Spatial and temporal patterns in a sagebrush steppe spider community (Arachnida: Araneae). J. Arachnol., 11: 31-50.
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
•
2. AITCHINSON, C. W. 1984. The phenology of winter-active spiders. J. Arachnol., 12: 249271.
•
3. ALBERT, R. 1976. Zusammensetzung und Vertikalverteilung der Spinnenfauna in
Buchenwäldern des Solling. Faun.-Ökol. Mitt., 5: 65-80.
•
4. ALBERT, R. 1982. Untersuchungen zur Struktur und Dynamik von Spinnengesellschaften
verschiedener Vegetationstypen im Hoch-Solling. HochschulSammlung Naturwissenschaft
Biologie Band 16. HochschulVerlag-Freiburg.
•
5. ALDERWEIRELDT, M. 1993. Fluctuations de l`activité saisonnière d`araignées dans les
champs de maïs et d`ivraie italien. Nwsbr. Belg. Arachnol. Ver., 8 (2): 32-43.
•
6. ALMQUIST, S. 1969. Seasonal growth of some dune-living spiders. Oikos, 20: 392-408.
•
7. ASEGINOLAZA, C.; GÓMEZ, D.; LIZAUR, X.; MONTSERRAT, G.; MORANTE, G.; SALAVERRIA, M. R. & URIBE-ECHEBARRIA, P. M. 1996. Vegetación de la Comunidad Autónoma del
País Vasco. Departamento de Ordenación del Territorio, Vivienda y Medio Ambiente.
Gobierno Vasco. 2ª edición. Vitoria. 361 pp.
•
8. ASCASO, C. & BARRIENTOS, J. A. 1986. Araneae: Comparación de los resultados anuales
de dos métodos de muestreo indirectos. Actas X Congreso Inter. Aracnol., Jaca, España. J.
A. Barrientos (Ed.). Vol. I: 175-182.
•
9. BAERT, L. L. & KEKENBOSCH, J. 1982. Araignées des Hautes Fagnes. II. Ecologie
(Ecologie of Belgian Spiders. III). Bull. Inst. R. Sci. Nat. Belg., 54 (1): 1-21.
•
10. BAERT, L. L.; KEKENBOSCH, J. & VANHERCKE, L. 1983. Araignées et opilions de la
Gaume dans les environs de la station biologique d`Ethe-Buzenol. Ecology of Belgian spiders IV. Bull. Inst. R. Sci. Nat. Belg., 55 (4) : 1-37.
•
11. BARRAQUETA, P. 1985. Zur Okologie von Pinus radiata-Forsten im Baskenland:
Streuproduktion, Streu-Abbau und Bodenmakrofauna. Dissertation, Universität Bremen, 1206.
•
12. BARRAQUETA, P. 1988. Las comunidades de diplópodos (Myriapoda) en diversos ecosistemas forestales del País Vasco. Biología Ambiental. II Congreso Mundial Vasco. Gobierno
Vasco. J.C. Iturrondobeitia (Ed.): 277-289.
•
13. BARRAQUETA, P. 2001. La fauna del suelo: entre la repoblación forestal y la tala. En:
Conservación, Uso y Gestión de los Sistemas Forestales. VI. Jornadas de Urdaibai sobre
Desarrollo Sostenible. M. Díez Salinas; M. C. de la Huerga & C. Giménez
(Coord.).Departamento de Ordenación del Territorio, Vivienda y Medio Ambiente. Gobierno
Vasco: 257-261.
•
14. BARRIENTOS, J. A. 1985a. Arañas, fenología reproductora y trampas de intercepción.
Actas II Congreso Ibér. Entomol. Sociedade Portuguesa de Entomologia. Suplemento I: 317326.
•
15. BARRIENTOS, J. A. 1985b. Artrópodos epígeos de San Juan de la Peña (Jaca, Huesca):
IX: Arañas agelénidas y hahnidas. Pirineos, 126: 81-131.
•
16. BARRIENTOS, J. A. 1985c. Artrópodos epígeos de San Juan de la Peña (Jaca, Huesca):
X: Arañas licósidas. Pirineos, 126: 133-162.
•
17, BARRIENTOS, J. A. & CARDOSO, P. 2007. The genus Malthonica Simon, 1898 in the
Iberian Peninsula (Araneae: Agelenidae). Zootaxa 1460: 59-68.
121
A. CASTRO GIL
•
18. BARRIENTOS, J. A.; ESPUNY, A. & ASCASO, C. 1994. Harpactea aeruginosa sp. n. y
Harpactea hombergi (Scopoli, 1763) (Araneae, Dysderidae) en el Montseny (Barcelona,
España). Eos, 69: 31-39.
•
19. BARRIENTOS, J. A.; ESPUNY, A. & ASCASO, C. 1996. Distribución espacial de los araneidos (Arachnida, Araneae) en un encinar montano del Montseny (Barcelona, España). Rev.
Suisse Zool. VHS: 29-44.
•
20. BARRIENTOS, J. A. & PUJADE, J. 1999. Nota sobre les aranyes de Santa Coloma
(Andorra) col·lectades amb trampa Malaise. Orsis, 14: 47-49.
• 21.
BLASCO, A. & RAMBLA, M. 1985. Artrópodos epígeos del macizo de San Juan de la Peña.
XII: Familias de araneidos de escasa representación. 5: Familias de arañas cribeladas.
Pirineos, 126: 221-227.
122
•
22. BONNET, P. 1935. Theridion tepidariorum C. L. Koch araignée cosmopolite. Répartitioncycle vital-moeurs. Bull. Soc. Hist. Nat. Toulouse, 68: 335-386.
•
23. BONTE, D.; MAELFAIT, J. P. & HOFFMANN, M. 2000. Seasonal and diurnal migration
patterns of the spider (Araneae) fauna of coastal grey dunes. Ekologia, 19 (supplement,4) :
5-16.
•
24. BOSMANS, R. 1995. Description de Bordea, nouveau genre endémique d`araignées des
Pyrénées (Araneae : Linyphiidae). Bull. Mus. Natl. Hist. Nat., Paris, 4e sér., 17 (1-2) : 87-94.
•
25. BOSMANS, R. 1997. Revision of the genus Zodarion Walckenaer, 1833, part II. Western
and Central Europe, including Italy (Araneae: Zodariidae). Bull. Br. Arachnol. Soc. 10 (8):
265-294.
•
26. BOSMANS, R. & CASTRO, A. 2002. Dos arañas nuevas para España (Araneae:
Theridiidae, Salticidae). Rev. Iber. Aracnol., 5: 51-53.
•
27. BOSMANS, R. & DE KEER, R. 1985. Catalogue des araignées des Pyrénées: especes citées, nouvelles recoltes et bibliographiques. Doc. Trav. Inst. r. Sci. Nat. Belg. 23: 1-68.
•
28. BRAUN, D. 1992. Aspekte der Vertikalverteilung von Spinnen (Araneae) an
Kiefernstämmen. Arachnol. Mitt., 4: 1-20.
•
29. BROEN, B. V. & MORITZ, M. 1963. Beiträge zur Kenntnis der Spinnentierfauna
Norddeutschlands. I. Über Reife- und Fortplanzungszeit der Spinnen (Araneae) und
Weberknechte (Opiliones) eines Moorgebietes bei Greifswald. Dtsch. Entomol. Z., 10 (III/V):
379-413.
•
30. BUDDLE, C. M. & DRANEY, M. L. 2004. Phenology of linyphiids in an old-growth decidoous forest in Central Alberta, Canada. J. Arachnol., 32: 221-230.
•
31. BUDDLE, C. M. & HAMMOND, H. E. 2003. Comparison of ground beetles (Coleoptera:
Carabidae) and spiders (Araneae) collected in pan and pitfall traps. Canadian Entomol., 135:
609-611.
•
32. CANARD, A. 1981. Utilisation comparée de quelques méthodes d`échantillonage pour
l`étude de la distribution des araignées en landes. Atti Soc. Tosc. Sci. Nat., Mem., ser. B, 88,
suppl. 84-94.
•
33. CANARD, A. 1982. Les araignées du Massif armoricain. II. Les Mimétidés. Bull. Soc. Sc.
Bretagne, 54 (1-6): 77-89.
•
34. CANARD, A. 1983. Les araignées du Massif armoricain. III. Les Atypidés. Bull. Soc. Sc.
Bretagne, 55 (1-4): 47-53.
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
•
35. CANARD, A. 1984. Contribution à la connaisance du développement, de l`écologie et de
l`écophysiologie des Aranéides de landes armoricaines. Thèse. Université de Rennes I.
•
36. CANARD, A. 2005. Catalogue of Spiders Species from Europe and the Mediterranean
basin. Parts I & II. Rev. Arachnol., 15 (3): 1-255.
•
37. CARDOSO, P. 2004. The use of Arachnids (Class Arachnida) in biodiversity evaluation
and monitoring of natural areas. Tese. Faculdade de Ciências. Universidade de Lisboa.
•
38. CARDOSO, P. 2008. Biodiversity and conservation of Iberian spiders: past, present and
future. Boln. S.E.A., 42 : 487-492.
•
39. CARDOSO, P. ; SILVA, I. ; DE OLIVEIRA, N. G. & SERRANO, A. R. M. 2007. Seasonality
of spiders (Araneae) in Mediterranean ecosystems and its implications in the optimum sampling period. Ecol. Entomol., 32 (5) : 516-526.
•
40. CARDOSO, P. ; SCHARFF, N. ; GASPAR, C. ; HENRIQUES, S. S. ; CARVALHO, R. ; CASTRO, P. H. ; SMICHDT, J. B. ; SILVA, I. ; SZÜTS, T. ; CASTRO, A. & CRESPO, L. C. 2008.
Rapid biodiversity assesment of spiders (Araneae) using semi-quantitative sampling : a case
study in a Mediterraneam forest. Insect Conservation and Diversity, 1 : 71-84.
•
41. CASTRO, A. 2004a. Catálogo preliminar de las arañas del País Vasco. En: Biodiversidad
y arácnidos: los invertebrados y la estrategia ambiental vasca de desarrollo sostenible. A.
Castro (Ed.). Munibe, suplemento 21: 44-69.
•
42. CASTRO, A. 2004b. Estudio biocenológico y faunístico del Orden Araneae (Arthropoda,
Arachnida) en los encinares cantábricos de Guipúzcoa y Navarra (España). Tesis Doctoral.
Universidad Autónoma de Madrid. Cd-Rom.
•
43. CASTRO, A. & ALBERDI, J. M. 2002. New spider species (Araneae) for the Spanish and
Iberian fauna found in the Basque Country (Northern Spain). Munibe (Ciencias NaturalesNatur Zientziak), 55: 175-182.
•
44. CATALÁN, P.; AIZPURU, I.; ARETA, P.; MENDIOLA, I.; DEL BARRIO, L. & ZORRAKIN, I.
1989. Guía ecológica de Artikutza (naturaleza y huella humana). Ayuntamiento de San
Sebastián. 103 pags.
•
45. CHRISTOPHE, T. & BLANDIN, P. 1977. The spider community in the litter of a coppiced
chesnut woodland (Forêt de Montmorency, Val d`Oise, France). Bull. Br. Arachnol. Soc., 4
(3): 132-140.
•
46. CHURCHILL, T. B. 1993. Effects of sampling methodology on the composition of a
Tasmanian coastal heathland spider community. Mem. Qld. Mus., 33: 475-481.
•
47. CHURCHILL, T. B. & ARTHUR, J. M. 1999. Measuring spider richness: effects of different
sampling methods and spacial and temporal scales. J. Insect Conserv., 3: 287-295.
•
48. CODDINGTON, J. A.; GRISWOLD, C.; DAVILA, D.; PEÑARANDA, E. & LARCHER, S.
1991. Designing and testing sampling protocols to estimate biodiversity in tropical systems.
In: The Unity of Evolutionary Biology, Vol.I. Proc. Fourth Int. Congress Syst. Evol. Biol. E.
Dudley (Ed.). Dioscorides Press, Portland, Oregon: 44-46.
• 49.
CODDINGTON, J. A.; YOUNG, L. H. & COYLE, F. A. 1996. Estimating spider species richness in a southern appalachian cove hardwood forest. J. Arachnol., 24: 111-128.
•
50. CURTIS, D. J. 1978. Community Parameters of the Ground Layer Araneid-Opilionid
Taxocene of a Scottish Island. Symp. Zool. Soc. Lond., 42: 149-159.
123
A. CASTRO GIL
•
51. CURTIS, D. J. 1980. Pitfalls in spider community studies (Arachnida, Araneae). J.
Arachnol., 8: 271-280.
•
52. CURTIS, D. J. & MORTON, E. 1974. Notes on spiders from tree trunks of different bark
texture; with indices of diversity and overlap. Bull. Br. Arachnol. Soc., 3 (1): 1-5.
•
53. DE BAKKER, D.; MAELFAIT, J. P.; DESENDER, K.; HENDRICKX, F. & DE VOS, B. 2002.
Regional variation in spider diversity of Flemish forest stands. In: European Arachnology,
2000. S. Toft & N. Scharff (Eds.). Aarhus University Press, Aarhus: 177-182.
•
54. DENIS, 1945. Notes sur les Érigonides X. Remarques sur le genre Entelecara E. Simon
avec la description de formes nouvelles du genre Plaesiocraerus E. Simon. Bull. Soc. Hist.
Nat. Toulouse, 80: 203-215.
•
55. DENIS, J. 1954. Araignées des environs d`Espingo (Haute-Garonne). Bull. Soc. Hist. Nat.
Toulouse, 89: 137-156.
•
56. DENIS, J. 1955. Recherches d`araignées dans les Pyrénées Centrales (de Barèges a
Gavarnie). Bull. Soc. Hist. Nat. Toulouse, 90: 142-156.
•
57. DENIS, J. 1965. Notes sur les Erigonides. XXVIII. Le genre Trichoncus (Araneae). Ann.
Soc. Ent. Fr. 1 (2): 425-477.
•
58. DEPARTAMENTO DE AGRICULTURA, PESCA Y ALIMETACIÓN. 2007.
Inventario Forestal CAE 2005. Gobierno Vasco. http://www.nasdap.ejgv.euskadi.net/r5015135/es/contenidos/informacion/inventario_forestal_index/es_dapa/inventario_forestal_index.html
•
59. DRESCO, E. 1957. Captures d`Araignées en Espagne (Campagnes biospeologiques de
1952 et 1954). Famille des Agelenidae. Speleon, Oviedo, VII (1-4): 119-124.
•
60. DRESCO, E. 1962. Araignées capturées en France dans des grottes ou des cavités souterraines. Annales de Spéléologie, XVII (1): 177-193.
• 61.
DRESCO, E. & HUBERT, M. 1971. Araneae speluncarum Hispaniae. I. Cuadernos de espeleología, Santander, 7 (5/6): 199-205.
• 62.
•
DUFFEY, E. 1956 Aerial dispersal in a known spider population. J. Anim. Ecol., 25: 85-111.
63. DUFFEY, E. 1969. The seasonal movements of Clubiona brevipes Blackwall and
Clubiona compta C. L. Koch on oak trees in Monks Wood, Huntingdonshire. Bull. Br.
Arachnol. Soc. 1 (3): 29-32.
• 64.
DUFFEY, E. 1972. Ecological survey and the arachnologist. Bull. Br. Arachnol. Soc., 2 (5):
69-82.
124
•
65. DUFFEY, E. 1993. A review of factors influencing the distribution of spiders with special
reference to Britain. Mem. Qld. Mus., 33 (2): 497-502.
•
66. EDGAR, W. D. 1971. The life-cycle, abundance and seasonal movement of the wolf spider, Lycosa (Pardosa) lugubris, in Central Scotland. J. Anim. Ecol., 40 (2): 303-322.
•
67. EDWARDS, C. A. & FLETCHER, K. E. 1971. A comparison of extraction methods for
terrestrial arthropods. In: Methods of Study in Quantitative Soil Ecology. J. Phillipson (Ed).
Blackwell Scientific Publications. Oxford and Edinburgh: 150-185.
•
68. FAGE, L. 1919. Etudes sur les araignées cavernicoles III: le genre Troglohyphantes. Arch.
Zool. Expér. Gén. 58: 55-148.
•
69. FAGE, L. 1931. Araneae, 5ª Série, précédée d`un essai sur l`evolution souterraine et son
déterminisme. Biospeologica LV. Arch. Zool. Expér. 71: 91-291.
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
•
70. FERRÁNDEZ, M. A. 1985a. Los Dysderidae (Araneae) de la fauna portuguesa. Actas II
Congreso Ibér. Entomol. Vol. IV: 17-24.
•
71. FERRÁNDEZ, M. A. 1985b. Artrópodos epígeos de San Juan de la Peña (Jaca, Huesca):
XIII: Familias de araneidos de escasa representación. 4: Familias de arañas oonópidas, segéstridas y disdéridas. Pirineos, 126: 219-221.
• 72.
FERRÁNDEZ, M. A. & ALBERDI, J. M. 2005. Nuevas citas de Mimétidos ibéricos (Araneae:
Mimetidae). La Tarántula, 3: 9-25.
•
73. FUJII, Y. 1998. Ecological studies on wolf spiders (Araneae: Lycosidae) in a northwest
area of Kanto Plain, Central Japan: Habitat preference observed by hand-sorting. Acta arachnol., 47 (1): 7-19.
•
74. GABBUT, P. D. 1956. The spiders of an oak wood in south-east Devon. Entomologist´s
mon. Mag., 92: 351-358.
•
75. GODALL, P. & PÉREZ, J. A. 1985. Artrópodos epígeos de San Juan de la Peña (Jaca,
Huesca): XIII: Familias de araneidos de escasa representación. 6: Arañas saltícidas. Pirineos,
126: 227-230.
•
76. GRIMM, U. 1986. Die Clubionidae Mitteleuropas: Corinninae und Liocraninae
(Arachnida, Araneae). Abh. naturw. Ver. Hamb. 27: 1-91.
•
77. HÄNGGI, A.; STÖCKLI, E. & NENTWIG, W. 1995. Habitats of Central European Spiders.
Centre Suisse de cartographie de la faune. Miscellania Faunistica Helvetiae, 4. 460 pp.
•
78. HARVEY, P. R.; NELLIST, D. R. & TELFER, M. G. (Eds.). 2002. Provisional Atlas of British
Spiders (Arachnida, Araneae). Volumes 1 & 2. Huntingdon: Biological Records Centre.
•
79. HAUGE, E. 2000. Spiders (Araneae) from square samples and pitfall traps in coastal
heathland, western Norway. Habitat preference, phenology and distribution. Fauna norv.,
20: 31-42.
•
80. HAUGE, E. & MIDTGAARD, F. 1986. Spiders (Araneae) in Malaise traps from two islands
in the Oslofjord, Norway. Fauna norv. Ser. B, 33: 98-102.
•
81. HEIMER, S. & NENTWIG, W. 1991. Spinnen Mitteleuropas. Paul Parey.
•
82. HEYDEMANN, B. 1960. Die biozönotische Entwicklung vom Vorland zum Koog. I. Teil:
Spinnen (Araneae). Abhandlugen der Mathemastisch-Naturwissenschaftlichen Klasse
Jahrgang. NR.11. Wiesbaden.
•
83. HIDALGO, I. L. 1986. Estudio de los Tomísidos de la Provincia de León (Araneae:
Thomisidae & Philodromidae). Diputación Provincial de León.
• 84.
HOLLANDER, J. den. 1971. Life histories of species in the Pardosa pullata group, a study
of ten populations in the Netherlands (Araneae, Lycosidae). Tijdschr. Entomol., 114 (8): 255281.
•
85. HÖREGOTT, H. 1960. Untersuchungen über die qualitative und quantitative
Zusammensetzung der Arthropodenfauna in der Kiefernkronen. Beitr. Entomol., 10: 891-916.
•
86. HORMIGA, G. & SCHARFF, N. 2005. Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus
Pecado. Zool. J. Linn. Soc., 143: 359-404.
•
87. HORTON, D. R.; MILICZKY, E. R.; BROERS, D. A.; LEWIS, R. R. & CALKINS, C. O. 2001.
Numbers, diversity and phenology of spiders (Araneae) overwintering in cardboard bands
125
A. CASTRO GIL
placed in pear and apple orchards of Central Washington. Ann. Entomol. Soc. Amer. 94 (3):
405-414.
•
88. HORVÁTH, R. & SZINETÁR, C. S. 1998. Study of bark-dwelling spiders (Araneae) on
black pine (Pinus nigra) I. Misc. Zool. Hung., 12: 77-83.
•
89. HORVÁTH, R. & SZINETÁR, C. 2002. Ecofaunistical study of bark-dwelling spiders
(Araneae) on black pine (Pinus nigra) in urban and forest habitats. Acta Biol. Debr., 24: 87-101.
•
90. HÖVEMEYER, K. & STIPPICH, G. 2000. Assesing spider community structure in a beech
forest: Effects of sampling method. Eur. J. Entomol., 97: 369-375.
•
91. HUHTA, V. 1965. Ecology of spiders in the soil and litter of finnish forests. Ann. Zool.
Fennici, 2: 260-308.
•
92. HUHTA, V. 1971. Succession in the spider communities of the forest floor after clear-cutting and prescribed burning. Ann. Zool. Fennici. 8: 483-542.
•
93. HUHTA, V. 1972. Efficiency of different dry funnels techniques in extracting arthropoda
from raw humus forest soil. Ann. Zool. Fennici. 9: 42-48.
•
94. HUHTA, V. & VIRAMO, J. 1979. Spiders active on snow in northern Finland. Ann. Zool.
Fennici, 16: 169-176.
•
95. ICONA. 1994. Segundo Inventario Forestal Nacional. 1986-1995. Comunidad Foral de
Navarra. Ministerio de Agricultura, Pesca y Alimentación. 304 pags.
• 96.
ITÄMIES, J. & ROUTSALAINEN, M. 1984. Phenology of wolf spiders (Araneae, Lycosidae)
at Hämeenkyrö, SW Finland in 1980. Memoranda Soc. Fauna Flora Fennica, 60: 145-152.
126
•
97. JENNINGS, D. T. & HILBURN, D. J. 1988. Spiders (Araneae) captured in Malaise traps in
spruce-fir forests of west-central Maine. J. Arachnol., 16: 85-94.
•
98. JERARDINO, M.; FERNÁNDEZ, J. L. & URONES, C. 1988. Activity of epigean spiders:
abundance and presence over time (forest ecosystems, province of Salamanca, Spain).
Biología Ambiental. Actas II Congreso Mundial Vasco. J. C. Iturrondobeitia (Ed.). Gobierno
Vasco: 351-370.
•
99. JOCQUÉ, R. 1973. The spider fauna of adjacent woodland areas with different humus
types. Biol. Jb. Dodonaea, 41: 153-178.
•
100. JUBERTHIE, C. 1954. Sur les cycles biologiques des araignées. Bull. Soc. Hist. Nat.
Toulouse, 89 (3-4): 209-318.
•
101. KEMPSON, D.; LLOYD, M. & GHELARDI, K. 1963. A new extractor for woodland litter.
Pedobiologia, 3: 1-21.
•
102. KIRCHNER, W. 1987. Behavioural and Physiological Adaptations to Cold. The
Biorhythms of Spiders. In: Ecophysiology of Spiders. W. Nentwig (Ed.). Springer-Verlag, New
York: 67-77.
•
103. KISS, B. & SAMU, F. 2002. Comparison of autumn and winter development of two wolf
spider species (Pardosa, Lycosidae, Araneae) having different life history patterns. J.
Arachnol., 30 (2): 409-415.
•
104. KNOFLACH, B. 1993. Das Männchen von Episinus theridioides SIMON (Arachnida:
Araneae, Theridiidae). Bull. Soc. Entomol. Suisse, 66 : 359-366.
•
105. KOOMEN, P. 1998. Winter activity of Anyphaena accentuata (Walckenaer, 1802)
(Araneae: Anyphaenidae). In: Proc. 17th Eur. Coll. Arachnol. P.A. Selden (Ed.): 223-225.
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
•
106. KOPONEN, S. 1976. Spider fauna (Araneae) of Kevo area, northermost Finland. Rep.
Kevo Subartic Res. Stat., 13: 48-62.
•
107. KOPONEN, S.; HAUKIOJA, E. & ISO-IIVARI, L. 1974. Comparison of spider catches and
weather in subartic conditions. Proc. 6th Int. Arach. Congress: 77-81.
•
108. KRAUS, O. & BAUR, H. 1974. Die Atypidae der West-Paläarktie Systematik. Verbreitung
und Biologie (Arach. Araneae). Abh. Verh. Naturw. Ver. Hamburg, 17: 85-116.
•
109. KRONESTEDT, T. 1968. Notes on the Swedish species of the genus Centromerus F.
Dahl (Araneae, Linyphiidae). A faunistic report with ecological remarks. Ent. Tidskr., 89:
111-127.
•
110. LANG, A. 2000. The pitfalls of pitfalls: a comparison of pitfall trap catches and absolute density estimates of epigeal invertebrate predators in arable land. J. Pest Science 73: 99106.
•
111. LE PERU, B. 2007. Catalogue et répartition des araignées de France. Rev. Arachnol., 16:
1-468.
•
112. LEDOUX, J. C. 1997-2000. Araignées des ripisilves du Rhône (Gard et Vauclause). Bull.
Soc. Et. Sci. Nat. Vauclause, 1997-2000 : 13-38.
•
113. LEDOUX, J. C.; EMERIT, M. & PINAULT, G. 1996. Les araignées et opilions de Nohèdes
(Pyrénées Orientales). Office pour l`information Eco-Entomologique du LanguedocRoussillon. 36 pp.
•
114. LENSING, J. R. ; TODD, S. & WISE, D. H. 2005. The impact of altered precipitation on
spatial stratification and activity-densities of springtails (Collembola) and spiders (Araneae).
Ecol. Entomol., 30 : 194-200.
•
115. LOGUNOV, D. V. 1992. A review of the spider genus Tmarus Simon, 1875 (Araneae,
Thomisidae) in the USSR fauna, with a description of new species. Siberian biol. J.,1: 6173.
•
116. LOIDI, J. & BASCONES, J. C. 1995. Memoria del Mapa de Series de Vegetación de
Navarra. Departamento de Ordenación del Territorio y Medio Ambiente. Gobierno de
Navarra. Pamplona. 99 pp.
•
117. LUCZAK, J. 1959. The community of spiders of the ground flora of pine forest. Ekol.
Pol., Ser. A, VIII (11): 285-315.
•
118. MAELFAIT, J. P.; BAERT, L.; BONTE, D.; DE BAKKER, D.; GURDEBEKE, S. & HENDRICKX, F. 2004. The use of spiders as indicators of habitat quality and anthropogenic disturbance in Flanders, Belgium. In: European Arachnology, 2002. F. Samu & C. Szinetár (Eds).
Plant Protection Institute & Berzsenyi College, Budapest: 129-141.
•
119. MANSOUR, F.; ROSEN, D. & SHULOV, A. 1980. Biology of the spider Chiracanthium
mildei (Arachnida: Clubionidae). Entomophaga, 25 (3): 237-248.
• 120.
MARC, P. 1990. Données sur le peuplement d`aranéides des troncs de pins. In: Comptes
rendus du XIIème Colloque européen d`Arachnologie, Paris. M. L. Celerier; J. Heurtault & C.
Rollard. (Eds.). Bull. Soc. eur. Arachnol. (NHS 1): 255-260.
•
121. MARC, P.; CANARD, A. & YSNEL, F. 1999. Spiders (Araneae) useful for pest limitation
and bioindication. Agric. Ecosyst. Environ., 74: 229-273.
•
122. MARTÍNEZ DE MURGUÍA, L. 2002. La taxocenosis de Hymenoptera en Artikutza
(Navarra). Bol. S.E.A., 31: 227-237.
127
A. CASTRO GIL
• 123.
MARTÍNEZ DE MURGUÍA, L.; VÁZQUEZ, M. A. & NIEVES-ALDREY, J. L. 2001. The familias of Hymenoptera (Insecta) in an heterogenous acidofilous forest in Artikutza (Navarra,
Spain). Frustula entomol., XXIV (XXXVII): 81-98.
•
124. MARTÍNEZ DE MURGUÍA, L.; VÁZQUEZ, M. A. & NIEVES-ALDREY, J. L. 2002.
Distributions of sawflies and aculeates in a heterogenous secondary acid forest in Artikutza
(Navarre) (Insecta: Hymenoptera). Munibe (Ciencias Naturales-Natur Zientziak), 53: 183204.
•
125. MELIC, A. 2000. Theridula gonygaster (Simon, 1873) en España (Araneae: Theridiidae).
Rev. Iber. Aracnol. 1: 37-38.
•
126. MELIC, A. 2001. Arañas endémicas de la península Ibérica e islas Baleares (Arachnida:
Araneae). Rev. Iber. Aracnol., 4: 35-92.
•
127. MÉNDEZ, M. 2003. Miscelanea Aracnológica. Rev. Iber. Aracnol. 7: 257-260.
• 128.
MERRET, P. 1968. The phenology of spiders on heathland in Dorset. Families Lycosidae,
Pisauridae, Agelenidae, Mimetidae, Theridiidae, Tetragnathidae, Argiopidae. J. Zool. London,
156: 239-256.
•
129. MERRET, P. 1969. The phenology of Lyniphiid spiders on heathland in Dorset. J. Zool.
London, 157: 289-307.
• 130.
MILLIDGE, A. F. 1975. Re-examination of the erigonine spiders “Micrargus herbigradus”
and “Pocadicnemis pumila” (Araneae: Linyphiidae). Bull. Br. Arachnol. Soc., 3 (6): 145-155.
128
•
131. MORANO, E. 2004. Introducción a la diversidad de las arañas iberobaleares. En:
Biodiversidad y arácnidos: los invertebrados y la estrategia ambiental vasca de desarrollo
sostenible. A. Castro (Ed.). Munibe, suplemento 21: 92-137.
•
132. MOULDER, B.C. & REICHLE, D. E. 1972. Significance of spider predation in the energy
dynamics of forest floor arthropod communities. Ecol. Monogr., 42: 473-498.
•
133. NENTWIG, W.; HÄNGGI, A.; KROPF, C. & BLICK, T. 2003. Spinnen Mitteleuropas
/Central European Spiders. An internet identification key. Version of 8.12.2003.
http://www.araneae.unibe.ch
•
134. NIEMELÄ, J.; PAJUNEN, T.; HAILA, Y.; PUNTTILA, P. & HALME, E. 1994. Seasonal activity of boreal forest-floor spiders. J. Arachnol., 22: 23-31.
•
135. NOFLATSCHER, M. T. 1993. Beiträge zur Spinnenfauna Südtirols – IV: Epigäische
Spinnen am Vinschgauer Sonnenberg (Arachnida: Aranei). Ver. nat.-med. Verein Innsbruck,
80: 273-294.
•
136. NORGAARD, E. 1951. On the ecology of two lycosid spiders (Pirata piraticus and
Lycosa pullata) from a Danish Sphagnum bog. Oikos, 3: 1-21.
•
137. PALMGREM, P. 1972. Studies on the spider populations of the surroundings of the
Tvärminne Zoological Station, Finland. Commentat. Biol., 52: 5-133.
•
138. PALMGREN, P. 1974a. Die Spinnenfauna Finnlands und Ostfennoskandiens. IV.
Argiopidae, Tetragnathidae und Mimetidae. Fauna Fennica, 24: 2-70.
•
139. PALMGREN, P. 1974b. Die Spinnenfauna Finnlands und Ostfennoskandiens. V.
Theridiidae und Nesticidae. Fauna Fennica, 26: 2-54.
•
140. PALMGREN, P. 1975. Die Spinnenfauna Finnlands und Ostfennoskandiens. VI.
Linyphiidae 1. Fauna Fennica, 28: 2-102.
SEASONAL
•
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
141. PALMGREN, P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII.
Linyphiidae 2. Fauna Fennica, 29: 2-126.
• 142.
PARKER, J. R. 1990. The life cycle of Theridion mystaceum L. Koch. Newsl. Br. Arachnol.
Soc., 58: 2-3.
•
143. PEARCE, J. L. & VENIER, A. L. 2006. The use of ground beetles (Coleoptera: Carabidae)
and Spiders (Araneae) as bioindicators of sustainable forest management: a review. Ecol.
Ind., 6: 780-793.
•
144. PEKÁR, S. 1999a. Life-histories and the description of developmental stages of
Theridion bimaculatum, T. impressum and T. varians (Araneae: Theridiidae). Acta Soc. Zool.
Bohem., 63: 301-309.
• 145.
PEKÁR, S. 1999b. Some observations on overwintering of spiders (Araneae) in two contrasting orchards in the Czech Republic. Agric. Ecosyst. Environ., 73: 205-210.
•
146. PESARINI, C. 1996. Note sur alcuni Erigonidae italiani, con descrizione di una nouva
specie (Araneae). Atti. Soc. it. Sci. nat. Museo civ. Stor. Nat. Milano, 135/1994 (II): 413-429.
•
147. PLATEN, R.; MORITZ, M. & BROEN, B. V. 1991. Liste der Webspinnen- und
Weberknechtarten (Arach.: Araneida, Opilionida) des Berliner Raumes und ihre Auwertung
für Naturschutzzwecke (Rote Liste). In: Rote Listen der gefärhrdeten Pflanzen und Tiere in
Berlin. A. Auhagen; R. Platen & H. Sukopp (Eds.). Landschaftsentwicklung und
Umweltforschung S 6: 169-205.
•
148. PLATNICK, N. I. 1999. Dimensions of biodiversity: targeting megadiverse groups. In:
The Living Planet in Crisis: Biodiversity Science and Policy. J. Cracraft & F. T. Grifo (Eds.).
Columbia University Press. New York: 33-52.
•
149. PLATNICK, N. I. 2008. The World Spider Catalog, version 8.5. Internet: http://research.amnh.org/entomology/spiders/catalog/index.html
•
150. PUJADE, J. 1996. Resultados preliminares obtenidos a partir de una trampa Malaise
situada en una zona mediterránea pirenaica. Pirineos, 147-148: 61-80.
•
151. RIBERA, C. 1980. Los Araneidos cavernícolas del País Vasco. Kobie, 10: 534-538.
•
152. RIBERA, C. & HORMIGA, G. 1985. Artrópodos epígeos de San Juan de la Peña (Jaca,
Huesca): XI. Arañas linífidas. Pirineos, 126: 163-210.
•
153. RICHTER, C. J. J.; HOLLANDER, J. den & VLIJM, L. 1971. Differences in breeding and
motility between Pardosa pullata (Clerck) and Pardosa prativaga (L. Koch), (Lycosidae,
Araneae) in relation to habitat. Oecologia, 6: 318-327.
•
154. RIECKEN, U. 1998. The importance of semi-natural landscape structures in an agricultural landscape as habitats for stenotopic spiders. In: Proc. 17th Eur. Coll. Arachnol. P. A.
Selden (Ed.). Edinburgh: 301-310.
•
155. RIVAS-MARTÍNEZ, S. 1994. Clasificación bioclimática de la Tierra. Fol. Bot. Matritensis,
11.
•
156. ROBERTS, M. J. 1985. The spiders of Great Britain and Ireland. Vol. I: AtypidaeTheridiosamatidae. Vol. II: Linyphiidae. Harley Books, Colchester.
•
157. ROBERTS, M. J. 1987. The spiders of Great Britain and Ireland. Vol. III: Colour plates.
Harley Books, Colchester.
•
158. ROBERTS, M. J. 1995. Spiders of Great Britain & Northern Europe. Collins Field Guide.
HarperCollins Publishers. London.
129
A. CASTRO GIL
• 159.
ROMANO, R. & FERRÁNDEZ, M. A. 1983. Dysdera scabricula Simon, 1882, nueva especie para la Península Ibérica con notas acerca de los Dysderidos de la provincia de Navarra.
Actas I Congreso Ibér. Entomol. Universidad de León: 685-697.
160. RUSSELL–SMITH, A. & SWANN, P. 1972. The activity of spiders in coppiced chesnut
woodland in southern England. Bull. Br. Arachnol. Soc., 2 (6): 99-103.
• 161. RUZICKA, V. 1997. Spiders (Araneae) of dwarf Norway spruces. Proc. 16th Eur. Coll.
•
Arachnol.. pp. 281-287.
•
162. RUZICKA, V.; BOHAC, J. & MACEK, J. 1991. Invertebrate animals from hollow trees in
the Trebon basin. Acta Mus. mer., Ceske Budejovice-Sci. nat., 31: 33-46.
•
163. SAARISTO, M. 1974. Taxonomical analysis of Theonina cornix (Simon, 1881), the typespecies of the genus Theonina Simon, 1929 (Araneae, Linyphiidae). Ann. Zool. Fennici, 11:
240-243.
•
164. SCHAEFER, M. 1976. Experimentelle Untersuchungen zum Jahreszyklus und zur Überwinterung von Spinnen (Araneida). Zool. Jb. Syst., Bd. 103: 127-289.
• 165.
SCHAEFER, M. 1977. Winter ecology of spiders (Araneida). Z. angew. Entomol., 83: 113-
134.
• 166.
SCHAEFER, M. 1987. Life Cycles and Diapause. In: Ecophysiology of Spiders. W. Nentwig
(Ed.). Springer-Verlag, Berlín: 331-347.
•
167. SCHABERREITER, I. 1999. Bestandsaufnahme ausgewählter epigäischer
Arthropodengruppen in einem Föhrenwald auf dem Eichkogel (Mödling, Niederösterreich).
1. Araneae. Verh. Zool. Bot. Ges. Österreich, 136: 87-108.
•
168. SCHARFF, N.; CODDINGTON, J. A.; GRISWOLD, C. E.; HORMIGA, G. & BJORN, P. P.
2003. When to quit? Estimating spider species richness in a northern European deciduous
forest. J. Arachnol., 31: 246-273.
•
169. SCHENKEL, E. 1938. Spinentiere von der Iberischen Halbinsel, gesammelt von Prof. Dr.
O. Lundblad, 1935. Ark. Zool. 30 A (24): 1-29.
•
170. SCHNEIDER, K. & DUELLI, P. 1997. Fangeffizienz von Fenster- und Malaisefallen im
Vergleich. Mitt. Dtsch. Ges. Allg. Angew. Ent., 11: 843-846.
•
171. SECHTEROVÁ, E. 1992. On the biology of species of the genus Coelotes (Araneae,
Agelenidae) in Central European mountains. Acta Entomol. Bohemoslov., 89: 337-349.
•
172. SEGERS, H. & MAELFAIT, J. P. 1990. Field and laboratory observations on the life cycle
of Coelotes terrestris and C. inermis (Araneae: Agelenidae). Bull. Soc. Eur. Arachnol., 1 : 314324.
•
173. SERRA, A. & VIVES, E. 1979. Campanya bioespeleològica a Guipúzcoa. Recull de
Treballs Espeleologics SIS-7: 19-26.
•
174. SIMON, E. 1884. Les Arachnides de France. Roret, Paris 5 (2): 181-420.
•
175. SIMON, E. 1918. Descriptions de plusieurs espèces d`Arachnides récemment découvertes en France (quatrième note). Bull. Soc. Entomol. France, 1918: 152-155.
• 176.
SIMON, E. 1914-1937. Les Arachnides de France. Tome VI. Synopsis général et Catalogue
des espèces françaises de l`ordre des Araneae. Roret-Mulo. Paris.
• 177.
SIMON, U. 1991. Temporal species serie of web-spiders (Arachnida: Araneae) as a result
of pine tree-bark structure. Bull. Soc. neuchâtel. Sci. nat., 116 (1): 223-227.
130
SEASONAL
DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
•
178. SNAZELL, R. ; JONSSON, L. J. & STEWART, J. A. 1999. Neon robustus Lohmander
(Araneae: Salticidae), a fennoscandian spider found in Scotland and Ireland. Bull. Br.
Arachnol. Soc., 11 (6): 251-254.
•
179. SORENSEN, L. L.; CODDINGTON, J. A. & SCHARFF, N. 2002. Inventorying and estimating subcanopy spider diversity using semiquantitative sampling methods in an afromontane forest. Environ. Entomol. 31 (2): 319-330.
•
180. STANDEN, V. 2000. The adequacy of collecting techniques for estimating species richness of grassland invertebrates. J. Appl. Ecol., 37: 884-893.
•
181. THOMAS, H. 1999. Nemesia simoni Cambridge: une petite mygale discrète mais commune en Aquitaine. Bull. Soc. linn. Bordeaux, 27 (1): 33-40.
•
182. TOFT, S. 1976. Life histories of spiders in a danish beech wood. Nat. Jutlandica, 19: 540.
•
183. TOFT, S. 1978a. Phenology of some Danish Beech-wood spiders. Nat. Jutlandica, 20:
285-304.
•
184. TOFT, S. 1978b. The life-history of Achaearanea lunata (Cl.) in Denmark, with a note
on Theridion varians Hahn (Araneae: Theridiidae). Bull. Br. Arachnol. Soc. 4 (5): 197-203.
• 185.
TOPPING, C. J. & LUFF, M. L. 1995. Three factors affecting the pitfall trap catch of linyphiid spiders (Aranea: Linyphiidae). Bull. Br. Arachnol. Soc. 10: 35-38.
•
186. TOPPING, C. J. & SUNDERLAND, K. D. 1992. Limitations to the use of pitfall traps in
ecological studies exmplified by a study of spiders in a field of winter wheat. J. Appl. Ecol.,
29: 485-491.
•
187. TOWNES, H. 1972. A light-weight Malaise trap. Entomol. News, 83: 239-247.
•
188. TRETZEL, E. 1954. Reife und Fortpflanzungszeit bei Spinnen. Z. Morphol. Ökol. Tiere,
42: 634-691.
•
189. TREZEL, E. 1961. Biologie, Ökologie und Brutpflege von Coelotes terrestris (Wider)
(Araneae, Agelenidae). Teil I: Biologie und Ökologie. Z. Morph. U. Okol. Tiere, 49: 658-745.
•
190. TURNBULL, A. L. 1960. The Spider Population of a Stand of Oak (Quercus robur L.) in
Wytham Wood, Berks., England. Can. Entomol., 92: 110-124.
•
191. UETZ, G. W. 1975. Temporal and spatial variation in species diversity of wandering spiders (Araneae) in deciduous forest litter. Environ. Entomol., 4 (5): 719-724.
•
192. UETZ, G. W. 1979. The influence of variation in litter habitats on spider communities.
Oecologia, 40: 29-42.
•
193. UETZ, G. W. & UNZICKER, J. D. 1976. Pitfall trapping in ecological studies of wandering spiders. J. Arachnol. 3: 101-111.
•
194. URONES, C. 1985a. Aportaciones al conocimiento de la distribución de los Thomisidae
(Araneae) en la Península Ibérica. Bolm. Soc. Port. Ent., 3 (Suplemento 1): 449-458.
•
195. URONES, C. 1985b. Artrópodos epígeos de San Juan de la Peña (Jaca, Huesca): VII:
Arañas clubionoideas. Pirineos, 126: 43-60.
•
196. URONES, C. 1986. La familia Philodromidae (Araneae) en el centro-oeste de la
Península Ibérica. Boln. Asoc. Esp. Entom., 10: 231-244.
•
197. URONES, C. 1996. Catálogo y atlas de las arañas de la familia Anyphaenidae en la
península Ibérica e islas Baleares. Graellsia, 52: 73-80.
131
A. CASTRO GIL
• 198.
URONES, C. 2000. El género Diaea Thorell, 1869 (Araneae, Thomisidae) en la Península
Ibérica. Boln. Asoc. Esp. Ent., 24 (1-2): 85-96.
•
199. URONES, C. & GÓMEZ, J. 1985. Fenología de algunas especies del género Xysticus C.
L. Koch, 1835 (Araneae, Thomisidae) en la zona centro-occidental de España. Stud. Oecol.,
VI: 245-265.
•
200. URONES, C.; BARRIENTOS, J. A. & ESPUNY, A. 1995a. El género Anyphaena Sundevall,
1833 (Araneae: Anyphaenidae) en la Península Ibérica. Boln. Asoc. Esp. Ent., 19 (1-2): 109-131.
•
201. URONES, C.; JERARDINO, M. & BARRIENTOS, J. A. 1995b. Datos fenológicos de
Gnaphosidae (Araneae) capturados con trampas de caída en Salamanca (España). Rev.
Arachnol., 11 (5): 47-63.
•
202. VANCLAY, J. K. 2004. Indicator groups and faunal richness. FBMIS, 1: 105-113.
•
203. VECIN, P.; GOITI, U. & SALOÑA, M. 2002. Artrópodos nocturnos y crepusculares capturados mediante trampa Malaise en Bizkaia, con especial referencia al orden Diptera. Est.
Mus. Cienc. Nat. de Álava, 17: 159-169.
•
204. VANUYTVEN, H. 1991. A redescription of Phoroncidia paradoxa (LUCAS, 1846)
(Araneae: Theridiidae). Phegea, 19: 103-105.
•
205. VLIJM, L. & KESSLER-GESCHIERE, A. M. 1967. The phenology and habitat of Pardosa
monticola, P. nigriceps and P. pullata (Araneae, Lycosidae). J. Anim. Ecol., 36: 31-56.
•
206. WEISS, I. 1995. Spinnen und Weberknechte auf Baumstämmen im Nationalpark
Bayerischer Wald. In: Proc. 15th Eur. Coll. Arachnol.. V. Ruzicka (Ed.). Institute of
Entomology, Ceske Budejovice: 184-192.
•
207. WIEBES, J. T. 1959. The Lycosidae and Pisauridae (Araneae) of the Netherlands. Zool.
Verh., 42 : 1-78.
•
208. WIEHLE, H. 1964. Über Hyptiotes gerhardti WIEHLE (Arach., Araneae). Senck. Biol., 45
(1): 81-85.
•
209. WILLIAMS, P. H.; GASTON, K. J. & HUMPHRIES, C. J. 1997. Mapping biodiversity value
worldwide: combining higher-taxon richness from different groups. Proc. R. Soc. Lond. B.,
264: 141-148.
•
210. WORKMAN, C. 1977. Population density fluctuations of Trochosa terricola Thorell
(Araneae: Lycosidae). Ecol. Bull. (Stockholm), 25: 518-521.
•
211. WOZNY, M. 1992. Wplyw wilgotnosci podloza na zgrupowania pajaków oraz dynamica liczebnosci gatunków dominujacych borów sosnowych wgórz Ostrzeszowskich. Acta
Universitatis Wratislaviensis, 1124 (Prace Zoologiczne XXIII): 25-82.
• 212.
WUNDERLICH, J. 1982. Mitteleuropäische Spinnnen (Araneae) der Baumrinde. Zeitschr.
angew. Entomol., 94/1: 9-21.
•
213. YSNEL, F. & CANARD, A. 1986. Réflexions sur les cycles vitaux des Araignées européennes, l`exemple des espèces à toiles géometriques. Mém. Soc. r. Belge Ent., 33: 213-222.
•
214. YSNEL, F. & CANARD, A. 1990. Régulations saisonnières des cycles biologiques des
Araignées européennes. Les Colloques de l`INRA, 52. Dourdan (France): 73-78.
- Fecha de recepción/Date of reception: 23/05/2008
- Fecha de aceptación/ Date of acceptance: 26/06/2008
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Appendix I.- Breakdown of the catches made in the litter layer in Gorbea using the Kempson
method. Abbreviations: m = males, f = females. Numbers without letter = immatures.
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Appendix II.- Breakdown of the catches made using epigaeic pitfall traps in the forest of Artikutza.
Abbreviations: m = males, f = females. Numbers without letter = immatures.
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BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix II. Continuation.
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Appendix II. Continuation.
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BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix II. Continuation.
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Appendix III.- Breakdown of the catches made using Malaise traps in the forest of Artikutza.
Abbreviations: m = males, f = females. Numbers without letter = immatures.
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(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix III. Continuation.
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Appendix III. Continuation.
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DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix III. Continuation.
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Appendix III. Continuation.
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DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix III. Continuation.
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Appendix III. Continuation.
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DYNAMICS OF FOREST SPIDERS
(ARACHNIDA: ARANEAE) IN THE TEMPERATE ZONE OF THE
BASQUE COUNTRY AND NAVARRA (NORTHERN SPAIN)
Appendix III. Continuation.
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Appendix IV. Breakdown of the catches made using bark traps in the alder forest of Igara.
Abbreviations: m = males, f = females. Numbers without letter = immatures.
146